UNIVERSITY  FARM 


4^ 


MINNESOTA   BOTANICAL    STUDIES 


GEOLOGICAL  AND  NATURAL  HISTORY  SURVEY  OF  MINNESOTA 
CONWAY  MACMILLAN,  State  Botanist 


MINNESOTA 


VOL.  II 


REPORTS  OF   THE    SURVEY 
AND   BULLETIN   OF   THE   DEPARTMENT 

BOTANICAL    SERIES 


IV 


1898  - 1902 
MINNEAPOLIS,    MINNESOTA 


PRECS  OF 
LANCASTER,    PA. 


TABLE   OF    CONTENTS. 


I.     Contributions  to  a  knowledge  of  the  lichens  of  Min- 
nesota.— III.     The  rock  lichens  of  Taylors  Falls, 
Bruce  Fink        .......        I 

II.  A  method  of  determining  the  abundance  of  second- 
ary species,  Roscoe  Pound  and  Frederic  E. 
Clements  .  .  .  .  .  .  .  19 

III.  List  of  fresh-water  alga3  collected  in  Minnesota  dur- 

ing 1896  and  1897,  Josephine  E.  Tilden     .          .      25 

IV.  Corrections  and  additions  to  the  flora  of  Minnesota, 

A.  A,  Heller 30 

V.     New  and  interesting  species  from  New  Mexico,  A. 

A.  Heller 33 

VI.     Some  Musci  of  the  International  Boundary,  John 

M.  Holzinger    .......     36 

VII.     The  influence  of  gases  and  vapors  on  the  growth  of 

plants,  Emil  P.  Sandsten  .          .          .          -53 

VIII.     Seedlings  of  certain  woody  plants,  Francis  Ramaley    69 
IX.     Comparative  anatomy  of  hypocotyl  and  epicotyl  in 

woody  plants,  Francis  Ramaley         .          .  87 

X.     Contribution  to  the  life-history  of    Ruinex,   Bruce 

Fink 137 

XI.     Observations  on  Gigartina,  Mary  E.  Olson    .          .   154 
XII.     Seed  dissemination  and  distribution  of  Razoumofskya 

robusta  (Engelm.)  Kuntze,  D.  T.  MacDougal .    169 

XIII.  Observations  on  Constantinea,  E.  M.  Freeman      .    175 

XIV.  Extension  of  plant  ranges  in  the  upper  Minnesota 

valley,  L.  R.  Moyer  ...  .191 

XV.  List  of  Hepaticae  collected  along  the  international 
boundary  by  J.  M.  Holzinger,  1897,  Alexander 
W.  Evans  ...  •  193 

XVI.     Observations  on  Chlorochytrium,  E.  M.  Freeman.   195 
XVII.     Observations  on  Rhodymenia,  Frederic  K.  Butters.  205 
XVIII.     Contributions  to  a  knowledge  of  the  lichens  of  Min- 
nesota.— IV.     Lichens  of  the  Lake  Superior  re- 
gion, Bruce  Fink       .          .          .          •  -215 
XIX.     Contributions  to  a  knowledge  of  the  lichens  of  Min- 
nesota.— V.     Lichens  of  the  Minnesota  valley  and 
southwestern  Minnesota,  Bruce  Fink         .          .   277 


VI 


TABLE    OF    CONTENTS. 


417 


XX.     A  synonymic  conspectus  of  the  native  and  garden 

Aquilegias  of  North  America,  K.   C.  Davis         .  331 
XXI.     A  synonymic  conspectus  of  the  native  and  garden 

Aconitums  of  North  America,  K.   C.  Davis        .  345 
XXII.     A  contribution    to    the    knowledge  of    the  flora  of 

southeastern  Minnesota,    W.  A.    Wheeler     .          .   353 

XXIII.  The    seed    and    seedling    of    the    western    larkspur 

{Delphinium      occidentale      Wats.),      Francis 
Ramaley    ........ 

XXIV.  A  preliminary  list  of  Minnesota  Erysipheaa,  E.  M. 

Freeman    ........  423 

XXV.     Native  and  garden  Delphiniums  of  North  America, 

K.  C.  Davis      .         .         .          .         .          .         .  431 

XXVI.     Native  and  cultivated  Ranunculi  of  North  America 

and  segregated  genera,  K.   C.  Davis  .          .          .  459 
XXVII.     A  synonymic  conspectus  of  the  native  and  garden 

Thalictrums  of  North  America,  K.   C.  Davis      .  509 
XXVIII.     Some    preliminary    observations    on     Dictyophora 

ravenelii  Burt.,   C.  S.  Scofield    ....   525 

XXIX.     A  preliminary  list  of  Minnesota  Uredineaa,  E.  M. 

Freeman    ........   537 

XXX.     A  new  species  of  Alaria,  De  Alton  Saunders        .  561 
XXXI.     A  preliminary  list  of  Minnesota  Xylariacese,  F.  K. 

Butters      ........  563 

XXXII.     A  contribution  to  the  knowledge  of  the  flora  of  the 

Red  river  valley  in  Minnesota,    W.  A.    Wheeler      569 

XXXIII.  Observations  on  Gigartina  exasperata  Harv.,  H. 

B.  Humphrey   .          .          .          .          .          .          .601 

XXXIV.  Observations  on  the  algae  of  the  St.  Paul  city  water, 

M.   G.  Fanning         ......  609 

XXXV.     Notes    on    some    plants    of    Isle    Royale,     W.    A. 

Wheeler     .          .          .          .          .          .          .          .   619 

XXXVI.     Revegetation  of  Trestle  island,  D.  Lange       .          .621 
XXXVII.     Violet  rusts  of  North  America,  J.    C.  Arthur  and 

E.    W.  D.  Hoi-way 631 

XXXVIII.     Observations  on  the  embryogeny  of  Nelumbo,  H. 

L.  Lyon     ........   643 

XXXIX.     Contributions  to  a  knowledge  of  the  lichens  of  Min- 
nesota.— VI.    Lichens  of  northwestern  Minnesota, 
Bruce  Fink       .          .          .          .          .          .          -657 

XL.     Corallinae  verae  of  Port  Renfrew,  K.  Yendo    .          .   711 
XLI.     Observations  on  Pterygophora,  Conway  MacMillan  723 


LIST    OF   AUTHORS. 


ARTHUR,  J.  C.,  and  HOLWAY,  E.  W.  D.     Violet  rusts  of  North 

America      .         .          .          .          .          .          .          .          .          .631 

BUTTERS,  F.  K.     Observations  on  Rhodymenia          .          .          .   205 
BUTTERS,  F.  K.     A  preliminary  list  of  Minnesota  Xylariaceaa  .  563 
CLEMENTS,  F.  E.,  see  POUND,  R.,  and  CLEMENTS,  F.  E. 
DAVIS,  K.  C.     A   synonymic   conspectus  of  the  native  and  gar- 
garden  Aquilegias  of  North  America  .          .          .          -331 
DAVIS,  K.  C.     A  synonymic  conspectus  of  the  native  and  gar- 
den Aconitums  of  North  America  "               .          .          .          .  345 

DAVIS,  K.  C.     Native  and  garden  Delphiniums  of  North  Amer- 
ica     .  .431 
DAVIS,  K.  C.     Native  and  cultivated  Ranunculi  of  North  Amer- 
ica and  segregated  genera    .......   459 

DAVIS,  K.  C.     A   synonymic  conspectus  of  the  native  and  gar- 
den Thalictrums  of  North  America     .....   509 

EVANS,  A.  W.     List   of  Hepaticse   collected  along  the  interna- 
tional boundary  by  J.  M.  Holzinger,  1897  .          .          .    193 
FANNING,  MARY  G.     Observations  on   the  algae  of  the  St.  Paul 

city  water  .........   609 

FINK,  BRUCE.     Contributions  to  a  knowledge  of  the  lichens  of 

Minnesota. — III.   The  rock  lichens  of  Taylors  Falls     .          .        I 
FINK,  BRUCE.     Contributions  to  the  life  history  of  Rumex          .    137 
FINK.  BRUCE.     Contributions  to   a  knowledge  of  the  lichens  of 

Minnesota. — IV.    Lichens  of  the  Lake  Superior  region         .   215 
FINK,  BRUCE.     Contributions  to  a  knowledge  of  the  lichens  of 
Minnesota. — V.   Lichens  of   the  Minnesota  valley  and  south- 
western Minnesota       .          .          .          .          .          .          .          •   277 

FINK,  BRUCE.     Contributions  to  a  knowledge  of  the  lichens  of 

Minnesota. — VI.   Lichens  of  northwestern  Minnesota  .   657 

FREEMAN,  E.  M.     Observations  on  Constantinea      .         .          •    175 
FREEMAN,  E.  M.     Observations  on    Chlorochytrium  .          .    195 

FREEMAN,  E.  M.     A  preliminary  list  of  Minnesota  Erysipheaa.  423 
FREEMAN,  E.  M.     A   preliminary  list  of  Minnesota   Uredineas.  537 
HELLER,    A.    A.      Corrections   and    additions    to    the    flora    of 

Minnesota  .........     30 

HELLER,  A.  A.     New  and  interesting  species  from  New  Mexico.     33 
HOLWAY,  E.  W.  D.     See  ARTHUR,  J.  C.,  and  HOLWAY,  E.  W. 
D. 

vii 


Vlll  LIST    OF    AUTHORS. 

HOLZINGER,  J.  M.     Some  Musci  of  the  International  Boundary.     36 
HUMPHREY,   H.    B.      Observations   on    Gigartina    exasperata 

Harv 60 1 

LANGE,  D.     Revegetation  of  Trestle  island         .          .          .          .621 
LYON,  H.  L.     Observations  on  the  .embryogeny  of  Nelumbo        .  643 
MACDOUGAL,   D.   T.      Seed   dissemination    and   distribution   of 
MACMILLAN,  CONWAY.     Observations  on  Pterygophora    .          .   723 

Razoumofskya  robztsta  (Engelm.)  Kuntze  .          .          .169 

MOYER,  L.  R.     Extension  of  plant  ranges  in  the  upper  Minne- 
sota valley  .          .          .          .          .          .          .          •          .191 

OLSON,  MARY  E.     Observations  on  Gigartina  .          .          .   154 

POUND,  R.,  and  CLEMENTS,  F.   E.     A  method  of   determining 

the  abundance  of  secondary  species       .          .          .          .  19 

RAMALEY,  FRANCIS.     Seedlings  of  certain  woody  plants   .          .     69 
RAMALEY,  FRANCIS.      Comparative  anatomy  of  hypocotyl   and 

epicotyl  in  woody  plants      .......     87 

RAMALEY,   FRANCIS.      The   seed   and    seedling  of    the  western 

larkspur  (Delphinium  occidentale)  Wats.  .          .          .   417 

SANDSTEN,  EMIL  P.     The  influence  of  gases  and  vapors  on  the 

growth  of  plants          ........     53 

SAUNDERS,  DE  ALTON.     A  new  species  of  Alaria     .         .          .561 
SCOFIELD,   C.   S.     Some    preliminary    observations    on   Dictyo- 

phora  ravenelii  Burt.  .          .          .          .          .          .          •   525 

TILDEN,  JOSEPHINE  E.     List  of  fresh-water  algae  collected  in 

Minnesota  during  1896  and  1897  •          •          •          •          •      25 

WHEELER,  W.  A.     A  contribution  to  the  knowledge  of  the  flora 

of  southeastern  Minnesota  .......  353 

WHEELER,  W.  A.     A  contribution  to  the  knowledge  of  the  flora 

of  the  Red  river  valley  in  Minnesota  .....  569 
WHEELER,  W.  A.  Notes  on  some  plants  of  Isle  Royale  .  .619 
YKNDO,  K.  Coralline  verae  of  Port  Renfrew  .  .  .  .71 


LIST   OF    PLATES. 


I. 

II. 
III. 
IV.J 

V. 

VI. 

VII. 

VIII. 

IX. 

X. 

XI. 

XII. 

XIII. 

XIV. 

XV. 

XVI. 

XVII. 

XVIII. 

XIX. 

XX. 

XXI. 

XXII. 

XXIII. 

XXIV. 

XXV. 

XXVI. 

XXVII. 

XXVIII. 

XXIX. 


RAMALEY,  FRANCIS. 
RAMALEY,  FRANCIS. 
RAMALEY,  FRANCIS. 
RAMALEY,  FRANCIS. 
RAMALEY,  FRANCIS. 
RAMALEY,  FRANCIS. 
RAMALEY,  FRANCIS. 
RAMALEY,  FRANCIS. 
FINK,  BRUCE. 
FINK.  BRUCE. 


Seedlings  of  various  woody  plants 
Seedlings  of  various  woody  plants 
Seedlings  of  various  woody  plants 
Seedlings  of  various  woody  plants 
Anatomy  of  Hypocotyl. 
Anatomy  of  Hypocotyl. 
Anatomy  of  Hypocotyl. 
Anatomy  of  Hypocotyl. 
Embryogeny  of  Rumex, 
Embryogeny  of  Rumex. 


FINK,  BRUCE.     Embryogeny  of  Rumex. 
FINK,  BRUCE.     Embryogeny  of  Rumex. 
OLSON,  MARY  E.     Anatomy  of  Gigartina. 
OLSON,  MARY  E.     Anatomy  of  Gigartina. 
MAcDouGAL,  D.  T.     Trees  of  Pinus  ponder osa  affected 

with  parasitic  Razoumofskya. 
MAcDouGAL,  D.  T.     Plants  of  Razoumofskya. 
FREEMAN,  E.  M.     Anatomy  of  Constantinea. 
Anatomy  of  Constantinea. 
Anatomy  of  Chlorochytrium. 
Anatomy  of  Rhodymenia. 

Vegetation  of  Southeastern  Minno 


Vegetation  of  Southeastern  Minne- 
Vegetation  of  Southeastern  Minne- 
Vegetation  of  Southeastern  Minne- 


FREEMAN,  E.  M. 

FREEMAN,  E.  M. 

BUTTERS,  F.  K. 

WHEELER,  W.  A. 
sota. 

WHEELER,  W.  A. 
sota. 

WHEELER,  W.  A. 
sota. 

WHEELER,  W.  A. 
sota. 

WHEELER,  W.  A.     Vegetation  of  Southeastern  Minne- 
sota. 

WHEELER,  W.  A.     Vegetation  of  Southeastern  Minne- 
sota. 

WHEELER,  W.  A.     Vegetation  of  Southeastern  Minne- 
sota. 

RAMALEY,  FRANCIS.     Seed  and  seedling  of  the  Western 
Larkspur. 

SCOFIELD,  C.  S.     Field  of  Dictyo-phora  ravenelti. 


LIST    OF    PLATES. 


XXX. 

XXXI. 

XXXII. 

XXXIII. 

XXXIV. 

XXXV. 

XXXVI. 

XXXVII. 

XXXVIII. 

XXXIX. 

XL. 

XLI. 

XLII. 

XLIII. 

XLIV. 

XLV. 

XLVI. 

XLVII. 

XLVIII. 

XLIX. 

L. 

LI. 

LII. 

LIII. 

LIV. 

LV. 

LVI. 

LVII. 

LVIII. 

LIX. 

LX. 

LXI. 

LXII. 


SCOFIELD,   C.    S. 
SCOFIELD,   C.    S. 

FREEMAN,  E.  M. 
SAUNDERS,  D.  A. 
WHEELER,  W.  A. 
WHEELER,  W.  A. 
WHEELER,  W.  A. 
WTHEELER,  W.  A. 
WHEELER,  W.  A. 
WHEELER,  W.  A. 
WHEELER,  W.  A. 
WHEELER,  W.  A. 
HUMPHREY,  H.  B. 
FANNING,  M.  G. 

water. 
FANNING,  M.  G. 

water. 

FANNING,  M.  G. 
G. 


Plant  of  Dictyophora  ravenelii. 
Anatomy  of  Dictyophora  ravenelii. 
Witches  Broom  on  Pinus  strobus. 
Alaria  curtipes. 

Vegetation  of  the  Red  river  valley. 

Vegetation  of  the  Red  river  valley. 

Vegetation  of  the  Red  river  valley. 

Vegetation  of  the  Red  river  valley. 

Vegetation  of  the  Red  river  valley. 

Vegetation  of  the  Red  river  valley. 

Vegetation  of  the  Red  river  valley. 

Vegetation  of  the  Red  river  valley. 

Anatomy  of  Gigartina. 
Chart  of  algal  forms  in  St.  Paul  city 

Chart  of  algal  forms  in  St.  Paul  city 


Forms  of  water  supply  algae. 
Forms  of  water  supply  algee. 

W.  D.     Spores  of 


FANNING,  M 

ARTHUR,  J.    C.,  and  HOLWAY,  E. 

violet  rust. 

LYON,  H.  L.     Embryogeny  of  Nelumbo. 
LYON,  H.  L.     Embryogeny  of  Nelumbo. 
LYON,  H.  L.     Embryogeny  of  Nelumbo. 
YENDO,  K.     Amphiroa. 
YENDO,  K.      Cheilosporum. 
YENDO,  K.      Cheilosporum. 
YENDO,  K.      Cheilosporum  and  Corallina. 
YENDO,  K.      Corallina. 
YENDO,  K.     Anatomy  of  Coral linoe  verag. 
MAcMiLLAN,  CONWAY,  Pterygophora  californica. 
MACMILLAN,  CONWAY,  Pterygophora  californica. 
MAcMiLLAN,  CONWAY.      Pterygophora  californica. 
MAcMiLLAN,  CONWAY.      Pterygophora  californica. 
MAcMiLLAN,  CONWAY.     Anatomy  of  Pterygophora. 
MACMILLAN,  CONWAY.     Anatomy  of  Pterygophora. 


DATES  OF  PUBLICATION  OF  THE  PARTS. 


PART    V. 

PART  VI. 


PART      I.     Pp.       i-  68  June  15,  1898. 

PART    II.     Pp.     69-194,  PI.  I.-XVIII Feb.  22,  1899. 

PART  III.     Pp.  195-352,  PL  XIX.,  XX Dec.  29,  1899. 

PART  IV.  Pp.  353-536,  PI.  XXI. -XXXI Aug.  15,  1900. 

Pp-  537-656,  P1.XXXII.-L Jjftrf*  I9°I- 

Pp.  657-780,  PI.  LI-LXII Mar.  i«f,  1902. 


XI 


I.    CONTRIBUTIONS   TO   A   KNOWLEDGE   OF   THE 

LICHENS   OF   MINNESOTA— III.   THE   ROCK 

LICHENS    OF   TAYLORS    FALLS. 


BRUCE  FINK. 


THE  COMPOSITION  AND  ORIGIN  OF  THE  FLORA. 

The  lichens  listed  in  this  paper  were  collected  during  two  reg- 
ular annual  excursions  of  the  Summer  School  of  the  University 
of  Minnesota.  The  date  of  collection  of  all  numbers  up  to  92 
is  August  15,  1896,  and  the  plants  were  collected  by  Professor 
Conway  MacMillan  and  the  writer,  on  the  Algonkian  igneous 
rocks  of  the  Interstate  Park,  or  on  earth  or  branches  in  the 
crevices  of  the  rocks.  On  the  I4th  of  August,  1897,  I  accom- 
panied another  excursion  to  examine  the  lichens  growing  on  the 
Cambrian  sandstone  exposures  near  the  park.  Numbers  93  to 
119  were  collected  by  me  during  this  second  trip,  and  more  spe- 
cies were  noted  on  the  sandstone,  which  had  been  collected  the 
year  before  on  the  igneous  rocks.  In  all,  24  lichens  were  found 
growing  on  both  sandstone  and  igneous  rocks,  22  on  the  igne- 
ous rocks  only,  20  on  the  sandstone  only,  10  on  earth  in  the 
crevices  of  the  igneous  rocks  and  2  on  roots  or  branches  in  the 
crevices.  The  last  lichen  of  the  list  was  found  on  old  boards 
and  is  recorded  here  because  rare  or  difficult  to  detect  and  new 
to  the  State. 

It  was  my  intention  when  I  went  to  the  park  to  publish  what- 
ever might  be  found  of  interest  with  the  last  number  of  this  ser- 
ies of  papers.  But  after  observing  the  field  it  became  apparent 
that  the  locality  is  one  of  great  interest  both  as  to  origin  and 
present  composition  of  its  lichen  flora  and  that  these  character- 
istic floral  features  could  be  presented  best  in  a  separate  paper. 
A  little  observation  showed  that  the  tree  lichens  do  not  differ  to 
any  noticeable  extent  from  those  about  Minneapolis,  and  I  con- 
sequently confined  my  collecting  to  the  igneous  rocks  and 


MINNESOTA    BOTANICAL    STUDIES. 


sandstone.  There  are  rocks  near  the  park  containing  lime,  but 
they  did  not  seem  to  support  any  lichens  of  special  interest. 

The  collecting  was  all  done  on  the  Minnesota  side  of  the 
river  for  the  reason  that  the  erosion  of  the  valley  has  been  such 
as  to  leave  better  exposures  of  rock  here  than  on  the  Wisconsin 
side.  I  had  collected  from  excellent  exposures  of  igneous  rocks 
in  New  England  and  have  since  visited  similar  ones  in  various 
parts  of  northern  Minnesota ;  but  I  have  never  seen  any  other 
equally  limited  area  of  rock  exposure  that  aroused  so  much  in- 
terest, because  of  richness  of  lichen  flora  and  evidence  of  mi- 
grations and  struggle,  as  did  this  little  area,  set  aside  for  an 
interstate  park.  During  the  trip  of  1897  I  noticed  that  the  con- 
stant tramp  of  feet  had  begun  to  kill  out  the  lichens  in  many 
places  so  that  the  impression  of  richness  is  beginning  to  fade, 
and  the  botanist  must  soon  seek  some  place  near  by,  if  such  ex- 
ists, where  he  may  study  this  rich  flora  in  its  natural  beauty. 

Professor  E.  E.  Edwards,  of  Lancaster,  Wisconsin,  writes 
thus  of  the  lichens  found  in  the  park:  "The  rocks  of  the 
Dalles  owe  their  beauty  and  variegated  tints  not  alone  to  the 
metal  oxides,  or  to  the  feldspar  or  hornblende  chiefly  composing 
them,  but  to  the  growth  of  minute  lichens  upon  their  surfaces, 
and  these  vary  in  color  according  to  the  dryness  or  moisture  of 
the  atmosphere.  We  have,  therefore,  in  these,  through  sunshine 
and  shadow  and  the  varying  seasons,  an  endless  and  almost 
kaleidoscopic  play  of  colors  that  makes  them  alike  the  delight 
and  despair  of  the  artist."  The  little  area,  being  one  of  great 
natural  beauty  and  set  apart  for  an  interstate  park,  will  always 
attract  thousands  of  visitors  annually  and  I  hope  to  present  in 
this  paper  thoughts  which  will  enable  the  botanist  who  has  a 
fair  knowledge  of  lichen  species  and  their  distribution  to  see  in 
this  wonderful  lichen  population  something  of  far  greater  inter- 
est than  mere  beauty. 

Some  comparisons  between  the  locality  now  under  considera- 
tion and  others  will  best  show  its  richness  in  rock  lichens.  The 
area  examined  covers  only  a  few  acres  of  surface  and  gave  66 
lichens  growing  on  rocks  as  a  result  of  two  days'  collecting. 
The  whole  region  about  Minneapolis  when  more  thoroughly 
worked  only  furnished  30  saxicolous  lichens,  and  the  whole  of 
Fayette  county,  Iowa,  only  50.  The  latter  region  is  surely 
better  than  the  average  for  rock  lichens,  and  I  have  studied  it 
for  six  years.  Probably  however,  the  fact  that  I  have  not  looked 


Fink:    ROCK  LICIIKNS   01    TAVI.OKS   FALLS.  3 

so  closely  in  the  last  two  localities  for  lichens  not^strictly  char- 
acteristic of  rocks,  but  still  growing  on  them  occasionally, 
about  offsets  the  limited  time  spent  in  examining  the  area  form- 
ing the  basis  of  the  thoughts  here  presented,  so  that  the  numbers 
given  above,  save  for  difference  in  areas  covered  in  the  three 
instances,  may  still  be  taken  to  represent,  approximately,  rela- 
tive richness  in  rock  lichens.  But  again,  the  Taylors  Falls 
area,  with  one  possible  exception,  gave  me  more  such  lichens 
collected  in  the  two  days  than  are  recorded  in  any  State  list 
hitherto  published. 

The  igneous  rocks  at  Taylors  Falls  are  essentially  like  those 
which  outcrop  occasionally  between  this  place  and  Lake  Su- 
perior, and  the  fact  that  Taylors  Falls  is  practically  the  south- 
ern limit  of  outcrop  of  these  rocks  furnished  the  first  suggestion 
of  the  interest  involved  in  an  analysis  of  their  lichen-flora. 
Of  the  79  species  and  varieties  collected,  only  8  are  plants  not 
yet  found  further  south  in  Minnesota  or  Iowa.  These  8  I  have 
also  found  along  Lake  Superior,  and  they  are  species  not  com- 
monly occurring  further  south  except  at  high  elevations.  These 
are  Biatora  rufonigra  Tuck.,  Lecidea  alboccertilescens  Schasr., 
three  forms  of  Bucllia  petraa  (Flot.,  Koerb.)  Tuck.,  Umbili- 
caria  dillenii  Tuck.,  Nephroma  helveticum  Ach.  and  Ephebc 
solida  Born.,  all  forms  found  on  the  igneous  rocks  and  none  of 
them  on  the  sandstone.  Subtracting  the  8  species  and  varieties 
leaves  58  rock  lichens,  nearly  all  of  which  occur  in  other  por- 
tions of  the  southern  half  of  Minnesota,  but  not  all  on  rocks. 
Those  not  known  to  flourish  on  rocks  in  other  parts  of  the  State 
grow  on  them  here  under  unusually  favorable  conditions  to  be 
explained  below. 

What  has  already  been  stated,  especially  the  last  paragraph 
above,  merely  suggests  the  problems  of  interest  which  I  shall 
attempt  to  discuss  and  which  involve  a  knowledge  of  geological 
conditions  present  and  past,  as  well  as  acquaintance  with  lichen- 
species  and  their  distribution  in  general.  For  the  geological 
data  concerning  this  area,  I  have  relied  largely  on  the  researches 
of  Dr.  Charles  P.  Berkey,  who  has  recently  studied  the  region 
including  Taylors  Falls  in  detail,  and  who  is  therefore  especially 
able  to  give  the  information  needed  for  my  purpose.  The  ques- 
tions which  I  shall  consider  below  are  those  which  thrust  them- 
selves upon  me  as  I  observed  and  studied  this  extremely  in- 
teresting lichen-flora ;  and  though  the  area  is  a  small  one,  the 


4  MINNESOTA    BOTANICAL    STUDIES. 

questions  involved  are,  it  seems  to  me,  none  the  less  worthy  of 
consideration  when  we  notice  that  it  is  one  of  a  series  of  similar 
areas  where  certain  floral  elements  have  become  isolated  and 
gradually  killed  out  by  others. 

The  8  species  commonly  found  farther  north  have  evidently 
migrated  southward,  and  there  are  at  least  two  views  as  to  time 
and  method  or  cause  of  migration  worthy  of  consideration. 
First,  the  northern  species  might  perhaps  have  migrated  from 
Lake  Superior  along  the  exposures  of  igneous  rocks  extend- 
ing from  the  lake  to  four  or  five  miles  below  Taylors  Falls  in 
quite  recent  times,  long  after  the  retreat  of  the  last  glacier. 
Second,  they  seem  undoubtedly  to  be  the  remnant  of  a  flora 
driven  south,  doubtless  from  some  region  far  north  of  Lake 
Superior,  by  the  advancing  glaciers  and  left  stranded  on  favor- 
able substrata  at  Taylors  Falls  as  the  southern  extremity  of  a 
flora  migrating  south  before  the  glacier  or  more  probably  mi- 
grating north  on  the  return  of  post-glacial  climate  in  the  north 
temperate  zone. 

The  outcrops  of  the  igneous  rocks  between  Lake  Superior 
and  Taylors  Falls  are  not  frequent  enough  to  make  either  theory 
seem  very  plausible ;  but  the  second  is  reasonable  since,  under 
the  influence  of  slow  decrease  in  temperature  to  the  southward, 
migration  would  naturally  follow  increasingly  favorable  cli- 
matic conditions  in  that  direction  even  if  the  outcrops  were  not 
more  frequent  than  now.  However,  it  seems  probable  that  at 
the  time  of  the  first  glacial  advance  the  outcrops  were  much 
more  numerous  than  now.  Also,  the  rock  lichens,  now  found 
on  the  igneous  rocks  only,  doubtless  found  a  foothold  on  the 
sandstones  on  the  line  of  retreat  under  the  more  favorable  cli- 
matic conditions  of  glacial  times.  The  numerous  boulders  of 
the  same  igneous  rocks,  scattered  over  the  ground  by  the  gla- 
ciers could  help  in  the  advance  southward  of  some  of  the 
species  since  glacial  times,  but  hardly  of  those  seldom  or  never 
found  on  boulders,  as  the  Umbilicaria  listed.  On  the  whole,  it 
seems  doubtful  whether  a  single  one  of  the  8  northern  species 
has  migrated  southward  in  post-glacial  times  under  increasingly 
unfavorable  conditions  as  to  climate  and  substrata. 

As  the  remnant  of  a  lichen-flora  driven  south  by  glaciers, 
these  plants  must  either  have  been  stranded  during  a  late  gla- 
cial advance,  as  during  a  slight  advance  during,  or  more  prob- 
ably after,  the  Wisconsin  stage ;  or  more  probably  have  been 


Fink:    ROCK  LICHEXS  OK  TAYLORS  FALLS.  5 

driven  further  south  than  their  present  position  by  each  of  the 
earlier  stages,  or  ages  as  the  case  may  be,  and  retreated  with 
each  return  of  interglacial  conditions.  The  8  northern  species 
at  Taylors  Falls  are  thus  either  a  few  of  a  former  flora  which 
has  doubtless  partly  died  out  and  partly  migrated  northward,  or 
possibly  a  few  species  which  migrated  to  the  locality  from  the 
mountains  to  the  east  and  west  during  a  late  glacial  stage,  as 
stated  above,  or  even  after  the  final  retreat  of  the  ice.  Igneous 
rocks  are  not  supposed  to  have  been  exposed  over  the  region 
covered  by  the  glaciers  south  of  the  area  now  under  considera- 
tion at  the  time  of  the  first  glacial  advance,  but  sandstone  no 
doubt  outcropped  frequently  and  probably  further  south  than  the 
glaciers  extended.  For  a  long  period  after  each  glacial  retreat 
the  surface  was  no  doubt  thickly  strewn  with  rocks  left  by  the 
melting  ice,  and  these  rocks  would  furnish  abundant  substrata 
for  a  retreat  of  the  saxicolous  lichens  to  the  north.  These  same 
boulders,  now  largely  covered,  would  partly  remain  at  the  sur- 
face during  interglacial  conditions  and  furnish  sufficient  foot- 
hold for  the  organisms  to  remigrate  during  a  subsequent  ad- 
vance of  the  ice,  thus  taking  the  place  of  the  sandstone  where 
it  was  covered  by  previous  drift  deposits.  Thus  several  migra- 
tions, alternately  southward  and  northward,  probably  followed 
in  succession,  and  we  are  studying  the  last  stage  in  the  last 
northward  retreat  in  this  not  yet  completed  series.  Of  course, 
it  is  apparent  that  the  Umbilicaria  and  many  other  lichens  not 
now  found  at  Taylors  Falls  might  have  flourished  on  the  sand- 
stone and  later  on  the  boulders  at  a  time  when  the  climate  was 
more  favorable  for  northern  species  than  now,  at  and  south  of 
the  area  under  consideration,  both  as  to  temperature  and  mois- 
ture, and  that  they  could  have  migrated  readily  enough  with  the 
advances  and  retreats  of  glacial  conditions.  What  species  of 
the  rock  lichens  were  able  to  endure  these  cycles  of  migration 
and  what  were  killed  out  is  not  easy  to  conjecture.  However, 
it  seems  certain  that  the  region  was  left  barren  of  such  life  and 
repopulated  several  times,  and  it  is  extremely  probable  that 
enough  species  survived  the  migrations,  or  possibly  in  part 
flowed  in  from  the  east  or  west  as  stated  above,  to  give  an  arctic 
or  subarctic  flora  at  Taylors  Falls  for  a  time  after  the  close  of 
the  ice  age. 

Since  the  time  when  this  last  northern  lichen-flora  became  es- 
tablished in  the  region  about  Taylors  Falls,  there  has  been  a 


MINNESOTA    BOTANICAL    STUDIES. 


gradual  change  toward  a  lichen-flora  characteristic  of  the  north- 
ern United  States  at  the  present  time.  No  doubt  the  8  northern 
species  now  found  on  the  igneous  rocks  are  being  rapidly  re- 
placed by  the  more  numerous  species,  which  are  better  adapted 
to  present  climatic  conditions.  With  the  exception  of  the  Neph- 
roma,  the  8  species  persisting,  all  lichens  confined  wholly  to 
rocky  substrata,  or  essentially  so,  and  being  therefore  favorably 
situated  as  to  substrata,  have  persisted  longest  against  un- 
favorable climatic  conditions  and  the  onslaughts  of  the  species 
which  are  to-day  surely  replacing  them.  Nephroma,  which 
is  arboricolous  as  well  as  rupicolous,  furnished  only  a  few  small, 
sterile  specimens  clinging  to  mossy  rocks.  Ephebe  was  seen 
in  one  spot  only,  and,  so  far  as  I  could  ascertain,  Umbi- 
licaria  persists  only  in  a  few  cool,  damp  or  shaded  spots. 
The  three  species  named  above,,  not  closely  attached  to  the 
rocks,  would  naturally  succumb  to  unfavorable  conditions  sooner 
than  those  named  below,  and  all  of  the  three  former  are  sterile 
and  apparently  just  on  the  verge  of  extermination  in  the  locality. 
The  other  three  species  all  grow  closer  to  the  rocks  and  are 
all  abundantly  fertile.  Biatora  seems  to  be  rare  and  is  not 
strictly  crustaceous  as  are  the  other  two.  Buellia  is  the  most 
common  of  the  8  northern  species  and  is  one  of  two  that  would 
be  expected  to  persist  longest  because  of  its  strictly  crustaceous 
habit.  Lecidea  is  as  thoroughly  crustaceous,  but  not  so 
common  as  the  Buellia.  It  must  be  noted  that  the  latter  plant 
shows  locally  the  strong  tendency  to  vary  so  characteristic  of 
organisms  attempting  to  adapt  themselves  to  change  in  environ- 
ment. Doubtless  this  variation  has  aided  the  plant  somewhat 
in  succeeding  best  of  all  the  present  or  former  more  northern 
species  of  the  locality  against  adverse  climatic  conditions. 
Whether  or  not  the  three  forms  of  the  species  listed  arose  from 
one  in  this  locality  has  no  particular  bearing  in  the  matter  as 
could  be  easily  shown.  Also  the  distribution  of  the  three  forms 
is  so  little  known  that  knowledge  as  to  which  particular  form 
is  most  common  locally  would  not  show  whether  it  is  one  most 
commonly  persisting  in  temperate  regions  or  not. 

It  is  interesting  to  note  the  time  involved  in  the  establishment 
of  the  Arctic  flora  and  the  change  from  this  to  the  present 
essentially  temperate  flora.  According  to  Professor  N.  H. 
Winchell's  views  as  to  the  recession  of  St.  Anthony  falls,  the 
final  retreat  of  the  glaciers  from  the  region  occurred  about  8,000 


Fink:    ROCK  LICHENS  OF  TAYLORS  FALLS.  7 

years  ago.  Thus  it  seems  that  the  succeeding  8,000  years  must 
have  sufficed  for  the  establishment  of  a  more  or  less  rich  Arctic 
flora  and  the  gradual  change  to  present  floral  conditions.  The 
relative  times  involved  in  the  establishment  of  the  first  flora  and 
the  gradual  change  to  the  present  cannot  be  arrived  at,  since  the 
richness  of  the  first  cannot  be  known,  and  we  cannot  yet  be  sure 
that  a  portion  of  the  species  migrating  southward  were  not 
killed  out  in  some  portions  of  the  series  of  migrations,  so  that 
some  portion  of  the  northern  species  that  became  established  in 
the  locality  would  have  to  migrate  toward  the  center  of  the  con- 
tinent from  the  southward-extending  mountains  already  men- 
tioned. Light  on  this  last  supposition,  which  can  only  be  fully 
obtained,  it  seems  to  me  at  present,  by  a  study  of  the  lichen  flora 
of  the  British  possessions  far  to  the  north  of  Minnesota,  would 
be  extremely  interesting. 

The  absence  of  the  8  northern  lichens  from  the  sandstone 
may  be  easily  explained,  since  it  seems  that  the  present  sand- 
stone surfaces  exposed  between  Lake  Superior  and  Taylors 
Falls  are  largely  or  entirely  due  to  post-glacial  erosion.  If 
some  of  these  surfaces  are  admitted  to  be  as  old  as  the  time  of 
the  last  glacial  retreat,  doubtless  Arctic  species  grew  on  them  at 
some  time  subsequent  to  that  retreat.  If  this  be  true,  it  is  yet 
easy  to  account  for  the  failure  of  these  lichens  to  persist  on  the 
sandstones  as  well  as  on  the  igneous  rocks,  since  the  lichen-flora 
of  these  porous  and  easily  eroding  surfaces  must  be  a  compara- 
tively changeable  and  transient  one,  so  that  whatever  such 
species  once  inhabited  them  would  now  be  replaced  by  species 
more  characteristic  of  present  climatic  conditions.  After  the 
final  retreat  of  the  ice  and  the  change  to  present  conditions 
of  temperature  and  moisture  began,  the  rapidly  eroding  surfaces 
would  begin  to  lose  their  northern  species  and  be  resupplied  by 
those  at  hand  on  other  substrata  at  once,  while  those  on  igneous 
rocks  could  be  replaced,  mainly,  at  least,  only  by  a  fierce  and 
long-continued  struggle  between  the  Arctic  and  temperate  floral 
elements.  The  large  number  of  species  found  on  the  sand- 
stones is  at  first  surprising,  for  while  the  igneous  rocks  are  much 
richer  in  individual  lichens,  they  show  no  appreciable  advantage 
in  species.  The  softer  texture  of  the  sandstone,  which  caused 
the  more  rapid  destruction  of  the  species  growing  under  un- 
favorable climatic  conditions,  has  also  brought  this  condition. 
To  be  a  little  more  explicit,  while  on  account  of  their  rapid  ero- 


8  MINNESOTA    BOTANICAL    STUDIES. 

sion  a  large  number  of  individuals  cannot  become  established 
on  them  and  remain  long  enough  to  constitute  a  flora  rich  in  in- 
dividuals, yet  because  of  the  porous  character  of  the  sandstone 
more  of  the  species  characteristic  of  temperate  regions  have 
doubtless  already  become  established  on  them  in  one  place  or 
another  than  on  the  igneous  rocks. 

While  the  8  species  and  varieties  so  fully  treated  above  are  of 
special  interest  there  are  some  thoughts  concerning  the  other  70 
(excluding  the  last  one  listed)  that  must  not  be  lost  sight  of. 
As  to  distribution  they  are  a  heterogeneous  group,  30  of  them 
being  pretty  generally  distributed  over  the  United  States  and 
Canada,  24  being  limited  so  far  as  their  distribution  is  known 
to  the  territory  east  of  the  Rocky  mountains,  7  being  thus  far 
found  only  in  the  northern  United  States  and  Canada  and  5  oc- 
curring throughout  the  United  States.  The  North  American  dis- 
tribution of  the  last  4  here  considered  is  so  little  known  that 
nothing  can  be  definitely  stated  of  it.  Of  these  70  all  but  4  or 
5  occur  on  rocks  in  some  other  portion  of  North  America,  but 
only  15  are  strictly  rupicoline.  Of  the  other  55,  some,  though 
more  characteristic  of  rocks,  are  more  or  less  frequently  found 
growing  on  other  substrata ;  and  others  actually  prefer  other 
substrata  and  are  growing  on  rocks  here  under  unusually  favor- 
able conditions.  These  lichens,  like  the  others,  are  of  course  the 
descendants  of  a  race  that  has  migrated  several  times.  Nearly 
all  of  them  being  species  also  occurring  in  Europe,  it  is  certain 
that  they  were  represented  by  like  species  during  early  Tertiary 
times,  far  to  the  north  where  our  continent  was  then  connected 
with  the  Eastern  continent  on  both  sides.  The  coming  of  a 
cooler  climate  and  finally  of  glacial  conditions  inaugurated  the 
series  of  migrations.  Finally  after  the  last  retreat  of  the  glaciers 
began,  the  55  species,  because  of  their  adaptation  to  more  than 
one  substratum,  would  follow  the  retreat  more  surely  and  more 
rapidly  and  thus  more  certainly  and  sooner  reach  a  given 
locality  and  begin  to  replace  a  flora  growing  under  unfavorable 
conditions.  To  just  what  extent  the  arctic  flora  would  become 
established  before  these  species  would  come  in  and  begin  to  re- 
place it  can  not  be  stated  since  the  rate  of  retreat  of  the  glaciers 
relative  to  the  rate  of  migration  of  essentially  stationary  organ- 
isms is  not  known. 

In  the  second  paper  of  this  series,  I  accounted  for  the  com- 
parative scarcity  of  lichens  about  Minneapolis  by  dryness  of 


Fink:   ROCK  LICHENS  OF  TAYLORS  FALLS.  9 

climate  and  stated  that,  were  it  not  for  peculiarly  unfavorable 
conditions  as  to  rock-formations,  this  explanation  would  require 
a  larger  per  cent,  of  the  total  number  of  lichens  found  there  to 
occur  on  the  rocks  because  of  greater  amount  of  moisture  near 
the  ground.  The  annual  precipitation  at  Osceola  Mills  for  the 
last  six  years  has  been  31.271  inches,  while  at  St.  Paul  it  has 
been  28.997  inches.  The  former  place  being  only  seven  miles 
from  Taylors  Falls,  the  figures  may  be  taken  to  show  that 
the  precipitation  in  the  locality  now  considered  is  about  2.274 
inches  more  per  annum  than  that  at  Minneapolis.  Hence  we 
have  at  Taylors  Falls  essentially  the  same  conditions  as  to  mois- 
ture of  atmosphere  as  at  Minneapolis.  However  at  the  former 
place  we  have  the  extensive  rock-formations  necessary  for  the 
establishment  of  the  plants,  and  we  find  further  that  the  igneous 
rocks  are  favorably  situated  for  lichen  development  in  that  they 
lie  along  a  river  course  formerly  better  shaded  than  now  and 
where  moisture  has  been  abundant  in  spite  of  comparative  dry- 
ness  of  atmosphere.  Also  this  flora  was  doubtless  largely  es- 
tablished when  the  climate  was  not  so  dry  as  now  and  is  persist- 
ing against  conditions  less  favorable  than  formerly  existed. 
Moreover  the  8  persisting  northern  species  add  to  the  number 
strictly  characteristic  of  present  climate  and  give  the  locality  a 
further  advantage  over  Minneapolis  and  vicinity.  Doubtless 
study  of  the  whole  lichen-flora  about  Taylors  Falls  would  show 
that  between  30  and  50  per  cent,  of  the  lichens  grow  on  rocks 
as  compared  with  12  per  cent,  at  Minneapolis.  The  slight  ad- 
vantage in  annual  precipitation  of  moisture  for  the  former  lo- 
cality, of  course,  adds  slightly  to  the  relative  richness  in  rock 
lichens,  but  this  is  insignificant  as  a  cause  when  compared  with 
the  elements  considered  above. 

Another  objection  of  doubtful  value  to  the  first  proposition 
suggested  to  account  for  the  invasion  of  the  northern  rock-floral 
elements  is  that,  though  there  is  a  continuous  line  of  conifers 
from  Lake  Superior  to  Taylors  Falls,  the  northern  tree-inhab- 
iting lichens  are  wholly  absent  at  Taylors  Falls,  or  so  scarce 
as  to  escape  notice.  The  coniferous  trees  are  not  so  conspicu- 
ous a  part  of  the  flora  at  Taylors  Falls  as  in  Pine  county,  fifty 
miles  north.  No  doubt  at  least  a  part  of  the  tree  lichens  char- 
acteristic of  more  northern  regions,  and  now  almost  certainly  to 
be  found  in  Pine  county,  have  extended  down  to  this  location 
in  post-glacial  times,  as  the  conifers  are  abundant  from  Lake 


10  MINNESOTA    BOTANICAL    STUDIES. 

Superior  down  to  the  southern  part  of  this  county,  and  with 
substrata  abundant,  they  could  do  so  in  spite  of  unfavorable 
climatic  conditions.  They  have  apparently  failed  to  advance 
as  far  as  Taylors  Falls,  because  of  favorite  substrata  becoming 
somewhat  scarce,  and  an  increase  of  unfavorable  conditions  as 
to  temperature  and  precipitation.  I  am  aware  that  the  glaciers 
probably  retreated  slowly  enough  so  that  forests  could  spring 
up  and  furnish  substrata  for  the  retreat  of  species  driven  south 
in  glacial  times,  before  they  would  die  out  at  the  south  on  ac- 
count of  the  return  of  warm  climate,  and  that  whatever  northern 
tree  lichens  exist  in  the  pineries  fifty  miles  north,  could  be  ac- 
counted for,  wholly  or  in  part,  as  having  migrated  from  the 
south.  Yet  I  am  quite  convinced  that  there  has  been  a  circula- 
tion of  arboreous  lichen-floral  elements,  between  Lake  Superior 
and  Pine  county,  in  post-glacial  time,  which  has  not  extended 
to  Taylors  Falls,  to  any  easily  observable  extent  surely,  though 
conditions  as  to  substrata  are  much  more  favorable  for  such 
lichens  to  move  southward  from  the  lake  than  for  the  rock 
lichens. 

Not  a  single  species  of  northern  lichen  was  found  in  the  rock 
crevices  or  soil  studied.  I  have  noticed  how  in  regions  recently 
burned  the  soil  becomes  literally  covered  in  places  by  lichens 
of  various  genera  in  five  to  fifteen  years,  and  there  can  be  no 
doubt  that  earth  lichens  took  possession  of  the  glacial  drift  rap- 
idly after  the  retreat  of  the  ice  began.  However,  from  the  very 
fact  that  lichens  spring  up  rapidly  on  earth,  the  species  charac- 
teristic of  temperate  climate  would  the  more  quickly  take  pos- 
session of  the  present  limited  amount  of  soil  available  for  lichen 
growth,  and  whatever  additional  amount  that  was  available 
when  the  strife  began  between  arctic  -and  temperate  earth  lich- 
ens, and  the  more  rapidly  kill  out  the  northern  species  once  in- 
habiting the  drift. 

A  consideration  of  the  statements  made  in  the  last  two  para- 
graphs and  various  other  portions  of  this  paper  points  to  the 
conclusion  that  a  study  of  the  whole  lichen-flora  of  the  area 
between  Taylors  Falls  and  Lake  Superior  is  essential  to  a  bet- 
ter understanding  of  the  problems  herein  considered.  In  the 
next  paper  of  this  series,  in  which  I  shall  consider  the  lichen- 
flora  of  the  Lake  Superior  region,  I  shall  be  able  to  show  ad- 
ditional reasons  for  the  study  of  this  territory.  It  is  one  of 
rapid  transition  in  lichen-flora,  and  after  a  study  of  the  areas  to 


Fink:   ROCK  LICHENS  OF  TAYLORS  FALLS.  11 

the  north  and  south  of  it,  questions  of  extreme  interest  have 
been  suggested  to  me  which  can  only  be  solved  by  a  study  of 
this  flora. 

The  principal  conclusions  are  as  follows  : 

(1)  The  region  considered  in  this  paper  is  an  important  one 

for  the  study  of  lichen-flora  because  of  position,  and 
geological  relations  past  and  present. 

(2)  The  flora  considered  is  one  of  great  interest  as  to  origin 

and  present  composition  and  as  to  evidence  of  struggle 
between  flora  elements. 

(3)  The  present  lichen-flora  is  composed  of  arctic,  sub-arctic 

and  temperate  florae  elements  in  which  the  last  have  long 
since  gained  the  advantage  and  are  killing  out  the 
others. 

(4)  It  is  not  supposed  that  the  northern  species  migrated  south 

in  post-glacial  times,  but  rather  that  this  flora  is  one  that 
followed  the  last  retreat  of  the  glaciers  and  was  for  a 
time  essentially  arctic,  having  since  changed  to  its  pres- 
ent composition. 

(5)  Reasons  for  the  above  conclusions  are  as  follows  : 

(a)  Southward  migration  would  more  naturally  result 
from  the  decrease  of  temperature  to  the  south 
inaugurated  by  the  on-coming  of  a  glacial  cli- 
mate and  would  thus  go  on  even  though  suit- 
able substrata  might  be  somewhat  scarce. 

(£)  But  during  the  glacial  advances  rocky  substrata 
were  doubtless  more  numerous  than  now,  a 
condition  adding  to  the  ease  of  migration. 

(c)  Under  the  influence  of  increasingly  favorable  cli- 

matic conditions  to  the  south,  the  plants  would 
lake  more  easily  to  unfavorable  substrata  and 
migrate  more  readily  on  this  account  also. 

(d)  The  8  northern  lichens  are  all  but  one  essentially 

rock  lichens  and  are,  therefore,  the  ones  that 
would  be  expected  to  persist  longest. 

(6)  The  northern  floral  elements  considered  may  have  been 

driven  south  during  a  late  glacial  advance  not  extending 
quite  to  the  region,  but  more  probably  have  been  forced 
to  migrate  further  south  several  times  and  migrate  north 
as  many  times. 


12  MINNESOTA    BOTANICAL    STUDIES. 

(7)  The   migrating  plants  may  have  been  in  part  or  wholly 

killed  out  in  some  part  of  the  series  of  migrations  south- 
ward in  the  Mississippi  Valley,  so  that  the  present  north- 
ern floral  element  would  have  to  flow  in  from  the  moun- 
tains to  the  east  and  west,  but  more  probably  found 
sufficient  substrata  and  were  not  killed  out  in  the  south- 
ward migrations  in  the  valley. 

(8)  The  time  involved  in  the  change  from  arctic  and  sub-arctic 

to  temperate  flora  is  probably  about  8,000  years.  The  re- 
lative times  occupied  in  the  establishment  of  the  northern 
flora  and  the  change  to  the  temperature  one  can  not  be 
estimated  at  present. 

(9)  The  absence  of  the  northern  floral  elements  from  the  sand- 

stone is  due  partly  to  the  fact  that  many  of  the  surfaces 
of  sandstone  exposures  are  post-glacial.  If  some  are  as 
old  as  the  last  retreat  of  the  glaciers  from  the  region,  the 
absence  is  still  easily  explained  since  the  surfaces  are 
easily  eroded  and  porous  so  that  floral  changes  go  on 
rapidly  on  these  rocks. 

(10)  Lichens  are  not  individually  numerous  on  the  sandstones 

because  of  this  easily  eroding  nature  which  causes  rapid 
change  and  destruction ;  yet  a  large  number  of  species 
become  established  in  one  place  or  another  on  them  be- 
cause lichens  quickly  gain  a  foothold. 

(n)  The  other  70  lichens  of  the  rocks  are  not  so  character- 
istically rock  lichens  and  would  migrate  more  easily  and 
rapidly,  and  the  more  quickly  reach  a  locality  and  re- 
place an  established  flora  existing  under  unfavorable 
conditions,  because  not  confined  to  one  substratum. 

(12)  The  rock  lichen-flora  of  the  locality  is  extremely  rich  be- 

cause of  abundance  of  rocks,  location  in  a  river  valley 
where  shade  and  moisture  have  been  plentiful  and  geo- 
graphically where  the  8  northern  species  have  persisted 
to  increase  the  number  more  characteristic  of  present 
climatic  conditions. 

(13)  In   substantiation  of  the  method  used  to  account  for  the 

present  composition  of  the  flora,  I  have  attempted  to  show 
that  the  absence  of  tree  and  earth  lichens  from  the  local- 
ity tends  to  prove  its  correctness. 

In  view  of  work  already  done  at  Taylors  Falls  and  along 
Lake  Superior,  the  study  of  the  region  of  rapid  transition 
in  lichen-flora  between  is  greatly  to  be  desired. 


Fink:   ROCK  LICHENS  OF  TAYLORS  FALLS.  13 

I  am  under  great  obligations  to  Dr.  Charles  P.  Berkey  for  in- 
formation concerning  present  and  past  geological  conditions  of 
the  area  studied.  My  thanks  are  also  due  to  Professor  Conway 
MacMillan  for  data  concerning  the  distribution  of  the  Conifers 
between  Taylors  Falls  and  Lake  Superior  and  to  Mr.  Geo.  H. 
Hazzard,  of  Taylors  Falls,  for  the  figures  of  annual  precipita- 
tion of  moisture  in  the  vicinity  of  Taylors  Falls. 

LIST   OF   SPECIES   AND   VARIETIES. 

1.  Ramalina  calicaris  (L.)  FR.  var.  farinacea  SCH^ER. 
On  igneous  rocks  and  sandstone,  no.  83. 

2.  Usnea  barbata  (L.)  FR.  var.  florida  FR. 
On  igneous  rocks  and  sandstone,  no.  45. 

3.  Usnea  barbata  (L.)  FR.  var.  rubiginea  MICHX. 
On  sandstone,  no.  117- 

4.  Theloschistes  lychneus  (NvL.)  TUCK. 
On  igneous  rocks  and  sandstone,  no.  64. 

5.  Theloschistes  concolor  (DICKS.)  TUCK. 
On  sandstone,  no.  102. 

6.  Parmelia  perforata  (JACQ^)  ACH. 
On  igneous  rocks,  no.  71. 

Not  previously  reported  from  Minnesota. 

7.  Parmelia  crinita  ACH. 
On  igneous  rocks,  no.  66. 

8.  Parmelia  borreri  TURN. 
On  sandstone,  no.  in. 

9.  Parmelia  saxatilis  (L.)  FR. 

On  igneous  rocks  and  sandstone,  nos.  52  and  67. 

10.  Parmelia  olivacea  (L.)  ACH. 
On  igneous  rocks,  no.  60. 

u.  Parmelia  caperata  (L.)  ACH. 

On  igneous  rocks  and  sandstone,  no.  50. 

12.  Parmelia  conspersa  (EHRH.)  ACH. 
On  igneous  rocks  and  sandstone,  no.  49. 

13.  Physcia  speciosa  (WuLF.  ACH.)  NYL. 
On  igneous  rocks  and  sandstone,  no.  63. 


14  MINNESOTA    BOTANICAL    STUDIES. 

14.  Physcia  aquila  (Acn.)  NYL.  var.  detonsa  TUCK. 
On  igneous  rocks  and  sandstone,  nos.  42  and  86. 
Not  previously  reported  from  Minnesota. 

15.  Physcia  pulverulenta  (SCHREB.)  NYL. 
On  igneous  rocks  and  sandstone,  no.  73. 

16.  Physcia  stellaris  (L.)  TUCK. 
On  igneous  rocks,  no.  2. 

17.  Physcia  tribacia  (Acn.)  TUCK. 
On  igneous  rocks,  nos.  55  and  77. 

18.  Physcia  csesia  (HOFFM.)  NYL. 
On  igneous  rocks,  no.  30. 

19.  Physcia  obscura  (EHRH.)  NYL. 
On  igneous  rocks,  nos.  5  and  47. 

20.  Pyxine  sorediata  FR. 
On  igneous  rocks,  no.  48. 

21.  Umbilicaria  dillenii  TUCK. 
On  igneous  rocks,  no.  87. 

22.  Nephroma  helveticum  ACH. 
On  igneous  rocks,  no.  26. 

Not  previously  reported  from  Minnesota. 

23.  Pel tigera  pulverulenta  (TAYL.)  NYL. 
On  earth  among  igneous  rocks,  no.  15. 

24.  Peltigera  rufescens  (NECK.)  HOFFM. 
On  earth  among  igneous  rocks,  no.  17. 

25.  Peltigera  canina  (L.)  HOFFM. 

On  earth  among  igneous  rocks  and  on  sandstone,  nos.  16  and 
28. 

26.  Peltigera  canina  (L.)  HOFFM.  var.  spuria  ACH. 
On  sandstone,  no.  119. 

27.  Peltigera  canina  (L.)  HOFFM.  var.  sorediata  SCH^ER. 
On  sandstone,  no.  96. 

28.  Pannaria  languinosa  (Acn.)  KOERB. 
On  igneous  rocks  and  sandstone,  no.  20. 

29.  Pannaria  microphylla  (SEV.)  DELIS. 
On  igneous  rocks  and  sandstone,  no.  35. 
Not  previously  reported  from  Minnesota. 

30.  Ephebe  solida  BORN.  ( ?) 
On  igneous  rocks,  no.  59. 

The  specimens  were  sterile  and  must  be  regarded  as  uncer- 


Fink:    ROCK  LICHENS  OF  TAYLORS  FALLS.  15 

tain  as  to  species.     I  found  the  same  plant  in  three  or  four  lo- 
calities in  northern  Minnesota. 

Not  previously  reported »from  Minnesota. 

31.  Collema  pulposum  (BERNH.)  NYL. 
On  earth  among  igneous  rocks,  no.  23. 

32.  Collema  flaccidum  ACH. 
On  igneous  rocks,  no.  43. 

33.  Leptogium  tremelloides  (L.  FIL.)  FR. 
On  igneous  rocks,  nos.  18,  25,  29  and  61. 
Not  previously  reported  from  Minnesota. 

34.  Leptogium  chloromelum  (SEV.)  NYL. 
On  sandstone,  no.  109. 

Not  previously  reported  from  Minnesota. 

35.  Placodium  elegans  (LINK.)  DC. 

On  igneous  rocks  and  sandstone,  no/53- 

36.  Placodium  cinnibarinum  (Acn.)  ANZ. 
On  igneous  rocks,  no.  31  and  62. 

37.  Placodium  aurantiacum  (LIGHT)  NAEG.  and  HEPP. 
On  igneous  rocks  and  sandstones,  no.  57. 

A  form  with  scanty  thallus  and  biatorine  apothecia,  appear- 
ing much  like  the  next. 

38.  Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP.  var.  sid- 
eritis  TUCK. 

On  igneous  rocks,  nos.  38,  39  and  91. 

39.  Placodium  vitellinum  (EHRH.)  NAEG.  and  HEPP. 
On  igneous  rock  and  sandstone,  no.  3. 

40.  Lecanora  rubina  (ViLL.)  ACH. 

On  igneous  rocks  and  sandstone,  no.  51. 

41.  Lecanora  subfusca  (L.)  ACH. 

On  igneous  rocks  and  sandstone,  nos.  12  and  115. 

42.  Lecanora  subfusca  (L.)  ACH.  var.  coilocarpa,  ACH. 
On  sandstone,  no.  108. 

43.  Lecanora  atra  ( HUBS.)  ACH.  (?) 
On  igneous  rocks,  no.  4. 

Not  previously  reported  from  Minnesota. 

44.  Lecanora  hageni  ACH. 
On  sandstone,  no.  118. 

45.  Lecanora  varia  (EHRH.)  NYL. 

On  igneous  rocks  and  sandstone,  no.  68. 


16  MINNESOTA    BOTANICAL    STUDIES. 

46.  Lecanora  varia  (EHRH.)  NYL.  var.  symmicta,  ACH. 
On  sandstone,  no  103. 

47.  Lecanora  cinerea  (L.)  SOMMERF/ 

On  igneous  rocks  and  sandstone,  nos.  u,  19,  34  and  70. 

48.  Lecanora  cinerea  (L.)  SOMMERF.  var.  laevata,  FR. 
On  igneous  rocks,  no.  14. 

Not  previously  reported  from  Minnesota. 

49.  Lecanora  fuscata  (SCHRAD.)  TH.  FR. 

On  igneous  rocks  and  sandstone,  nos.  33,  56,  94,  and  106. 
Not  previously  reported  from  Minnesota. 

50.  Rinodina  sophodes  (Acn.)  NYL. 

On  igneous  rocks  and  sandstone,  nos.  59  and  105. 

51.  Pertusaria  velata  (TURN.)  NYL. 
On  sandstone,  no.  95. 

52.  Pertusaria  communis  DC. 
On  sandstone,  no.  112. 

53.  Urceolaria  scruposa  (L.)  NYL. 

On  igneous  rocks  and  sandstone,  no.  9. 

54.  Stereocaulon  condensatum  HOFFM. 
On  sandstone,  no.  93. 

Not  previously  reported  from    Minnesota.      Also   not  listed 
before  west  of  New  England. 

55.  Cladonia  mitrula  TUCK. 
On  sandstone,  no.  99. 

56.  Cladonia  cariosa  (Acn.)  SPRENG. 
On  sandstone,  114. 

57.  Cladonia  pyxidata  (L.)  FR. 

On  earth  among  igneous  rocks,  no.  65. 

58.  Cladonia  squamosa  HOFFM. 

On  earth  among  igneous  rocks,  nos.  72  and  78. 
Not  previously  reported  from  Minnesota. 

59.  Cladonia  caespiticia  (PERS.)  FL. 

On  earth  among  igneous  rocks  and  on  sandstone,  nos.  68a  anc 
101. 

60.  Cladonia  fimbriata  (L.)  FR.  var.  tubaeformis  FR. 
On  sandstone,  nos.  104  and  no. 

61.  Cladonia  gracilis  (L.)  NYL. 

On  earth  among  igneous  rocks  and  on  sandstone,  no.  74. 


Fink:   ROCK  LICHENS  OF  TAYLORS  FALLS.  17 

62.  Cladonia  gracilis  (L.)  NYL.  var.  verticillata,  FR. 
On  sandstone,  no.  116. 

63.  Cladonia  delicata  (EHRH.)  FR. 

On  old  wood  among  igneous  rocks,  nos.  79*  and  67. 
Not  previously  reported  from  Minnesota. 

64.  Cladonia  rangiferina  (L.)  HOFFM. 
On  earth  among  igneous  rocks,  no.  81. 

65.  Cladonia  rangiferina  (L.)  HOFFM.  var.  alpestris  L. 
On  earth  among  igneous  rocks,  no.  82. 

66.  Cladonia  macilenta  (EHRH.)  HOFFM. 

On  old  roots  among  igneous  rocks,  no.  79*"  and  So*. 

67.  Biatora  rufonigra  TUCK. 
On  igneous  rocks,  no.  i. 

Not  previously  reported  from  Minnesota. 

68.  Biatora  coarctata(SM.  NYL.)  TUCK.  var.  brajeriana  SCH^R. 
On  sandstone,  no.  92. 

Not  previously  reported  from  Minnesota. 

69.  Biatora  myriocarpoides  (NYL.)  TUCK. 

On  sandstone,  no.  100.  Habitat  unusual,  but  I  cannot  distin- 
guish sufficiently  between  this  and  my  wood  specimens  to  separate 
them. 

Not  previously  reported  from  Minnesota. 

70.  Lecidea  albocaerulescens  (WULF.)  SCH^R. 
On  igneous  rocks  and  sandstones,  no.  27. 
Not  previously  reported  from  Minnesota. 

71.  Buellia  spuria  ARN. 

On  igneous  rocks,  no.  22. 

Not  previously  reported  from  Minnesota. 

72.  Buellia  petraea  (FLOT.,  KOERB.)  TUCK. 
On  igneous  rocks,  no.  58. 

Not  previously  reported  from  Minnesota. 

73.  Buellia  petraea  (FLOT.,  KOERB.)  TUCK.  var.  montagnaei 
TUCK. 

On  igneous  rocks,  no.  89. 

Not  previously  reported  trom  Minnesota. 

74.  Buellia    petraea    (FLOT.,    KOERB.)    TUCK.    var.    grandis 
FLOERK. 


18  MINNESOTA    BOTANICAL    STUDIES. 

On  igneous  rocks,  no.  89*.     Thallus  coarser,  more  crowded 
and  lighter  colored.     Hypothallus  deficient. 
Not  previously  reported  from  Minnesota. 

75.  Endocarpon  miniatum  (L.)  SCH^ER. 
On  igneous  rocks  and  sandstone,  no.  41. 

76.  Endocarpon  hepaticum  HEDW. 
On  sandstone,  no.  97. 

77.  Verrucaria  fuscella  FR. 
On  igneous  rocks,  no.  21. 

78.  Verrucaria  muralis  ACH. 
On  sandstone,  no.  107. 

79.  Thelocarpon  prasinellum  NYL. 
On  old  boards,  no.  113. 

Not  previously  reported  from  Minnesota. 


II.     A   METHOD   OF   DETERMINING   THE  ABUN- 
DANCE  OF   SECONDARY   SPECIES. 


ROSCOE  POUND  AND  FREDERIC  E.  CLEMENTS. 


In  determining  the  abundance  of  species,  appearances  are 
extremely  deceptive.  One  who  has  worked  over  the  prairies 
for  many  seasons  comes  to  think  that  he  can  pick  out  instantly 
the  most  abundant  secondary  species.  Long  continued  obser- 
vation in  the  field  stamps  a  picture  on  one's  mind,  and  it  seems 
a  simple  matter  to  pick  out  the  several  species  and  to  classify 
them  in  the  several  grades  of  abundance  with  reasonable  ac- 
curacy. As  a  matter  of  fact,  this  is  not  possible.  After  more 
than  ten  years  of  active  field  work  on  the  prairies,  it  seemed  to 
the  writers  that  the  mental  pictures  acquired  was  approximately 
sufficient  to  make  the  reference  of  the  commoner  secondary 
species  of  prairie  formations  to  their  proper  grades  an  easy  task. 
When  actual  looking  at  the  prairies  as  the  season  permitted 
appeared  to  confirm  the  picture  already  formed,  this  seemed 
certain.  Closer  analysis  of  the  floral  covering  proved  that  the 
conclusions  formed  from  looking  at  the  prairie  formations  and 
from  long  field  experience,  without  actual  enumeration  of  indi- 
vidual plants,  were  largely  erroneous.  The  psoraleas,  prairie 
clovers  and  blazing  stars  would  probably  occur  to  all  as  among 
the  most  abundant  of  the  secondary  species  in  the  vernal,  estival 
and  serotinal  aspects  of  the  prairies  respectively.  When  we 
first  addressed  ourselves  to  the  task  of  assigning  to  each  of  the 
various  prairie  species  its  proper  degree  of  abundance,  it  oc- 
curred to  us  at  once  that  we  could  take  a  certain  species,  or 
certain  species,  as  types  for  each  grade,  and  use  these  species 
as  standards  by  which  to  measure  the  others.  It  proved  in  the 
end  that  the  species  selected,  though  of  the  commonest  occur- 
rence and  hence  familiar  from  daily  observation,  were  in  many 
cases  referred  to  wrong  grades  as  compared  with  other  species, 
no  less  common,  but  for  some  reason  not  so  prominent.  The 
difficulty  is  that  the  species  which  appear  most  prominent  in  the 
constitution  of  the  prairies  are  not  necessarily  the  most  abundant. 


20  MINNESOTA    BOTANICAL    STUDIES. 

The  prominent-flowered  blazing  stars  and  prairie  clovers  make 
a  much  greater  impression  on  the  eye  than  species  which  are  far 
more  abundant,  and  the  same  thing  is  true  to  a  less  degree  of 
many  other  species.  To  insure  accurate  or  even  approximately 
accurate  results,  it  is  necessary  to  resort  to  some  method  of  actual 
count. 

Actual  count  is  usually  practicable  only  when  copious,  gre- 
gario-copious  or  sparse  plants  are  in  question.  But  it  is  only 
with  respect  to  such  species,  which  are  as  a  rule  secondary  in 
formations,  that  it  is  important  to  determine  minutely  the  grade 
of  abundance  manifested. 

During  the  past  season,  in  order  to  determine  the  actual  quan- 
titative relations  of  the  copious  and  gregario-copious  species, 
we  have  made  a  large  number  of  enumerations  of  the  individual 
plants  of  each  secondary  species  present  in  plots  five  meters 
square  in  characteristic  formations  of  each  of  the  four  phytogeo- 
graphical  regions  represented  in  Nebraska.  The  plot  used,  five 
meters  square,  is  as  large  as  can  be  used  to  insure  accuracy  in 
counting.  The  deficiences  resulting  from  the  small  size  of  the 
plots  are  corrected  by  taking  a  large  number  of  plots  at  each 
station  and  averaging  the  results.  There  is  a  surprisingly  close 
agreement  in  figures  obtained  from  plots  in  widely  separated 
stations  in  the  same  district,  provided  reasonable  care  is  taken 
to  "locate  them  in  typical  situations. 

By  way  of  illustration,  a  number  of  observations  are  given  in 
full.  These  are  not  averages,  but  are  the  actual  counts  as  taken 
in  the  field.  The  two  immediately  following  were  taken  on  the 
prairie  14  miles  northeast  of  Lincoln  in  the  prairie  grass  forma- 
tion ( Sporobolus-Koeleria-Pamcum).  The  second  was  made 
about  400  yards  distant  from  the  first. 

Amorpka  canescens         ......  387 

Aster  multiflorus       .          .          .          .          .          .          .223 

Antennaria  campestris  (16  patches)       .          .          .  209 

Solidago  rupestris     .          .          .          .          .          .  101 

Helianthus  rigidus         .          .          .          .          .          .  97 

Kuhnistera  Candida            ......  43 

Kuhnistera  purpurea     .          .          .          .          .          .  31 

Brauneria  pallida     .......  24 

Solidago  rigida     .          .          .                     .          .          .  19 

Kuhnia  glutinosa      ....... 

Comandra  umbellata      .          .                    .          .          .  7 

Rosa  arkansana  2 


Pound  and  Clements:    METHOD  OF  DETERMINING  SPECIES.     21 

(2) 
Amorpha  cancsccns         ......          -268 

Aster  multijlorus       .......      -205 

Antennaria  campestris  (6  patches)          .          .          .          154 
SoUdago  rupestris     .          .          .          .          .          .          .105 

Ifelianthus  rigidus        ......  63 

Kuhnistcra  Candida  ......        26 

Kuhnistera  purpurea    .          .          .          .          .          .  22 

Brauneria   pallida  .          .          .          .          .          .          10 

Solidago    rigida  .          .          .          .          .          .          .16 

Rosa  arkansana        .......  8 

SoUdago  rigidiuscula  ......      5 

Gerardia  purpurea  ......  3 

Laciniaria  scariosa      .......       2 

Erlgcron  ramosus  ......  i 

Linum  rigidum  was  prominent,  but  did  not  occur  in  either  of 
the  plots,  and  in  comparatively  few  of  those  laid  out. 

In  a  large  number  of  plots,  Amorpha  canescens  averaged  309, 
Aster  multijlorus  275,  and  Antennaria  campestris  12  patches 
and  145  individuals. 

Enumerations  were  also  made  in  the  same  formation  in  the 
transition  area  between  the  prairie  region  and  the  sand  hill  re- 
gion. The  following  example  is  one  of  a  number  made  south 
of  Broken  Bow  (Custer  County).  The  formation  is  the  ordi- 
nary prairie  grass  {Sporobohis-Koeleria-Panicum)  formation, 
modified  somewhat  on  account  of  the  sandy  soil. 

Amorpha  canescens         .          ,          .          •          .          .  291 
Aster  multijlorus       .......      238 

Kuhnistera  Candida  (?)          .          .          .          .          .  23 

Solidago  rupestris     .          .          .          .          .          .  .21 

Brauneria  pallida           .          .          .          .          .          .  17 

Helianthus  rigidus  .          .          .          .          .          .  .12 

Kuhnia  glutinosa            ......  5 

The  marked  decrease  in  the  number  of  secondary  species  and 
in  the  abundance  of  each  is  characteristic  of  this  transition  area. 

Another  count,  made  where  the  prairie  grass  formation  was 
giving  way  to  the  buffalo  grass  formation  on  the  one  hand,  and 
to  the  bunch  grass  formation  on  the  other,  is  interesting. 
While  the  prairie  grasses  (species  of  Sporobolus,  Koeleria  cris- 
tata  and  Panicum  Scribnerianuni)  were  controlling,  there  was 
a  strong  admixture  of  Bouteloua  oligostachya,  and  two  bunches 


22  MINNESOTA    BOTANICAL    STUDIES. 

of  Andropogon  scopartus  occurred  in  the  plot.  The  locality 
was  about  six  miles  northwest  of  the  preceding. 

Amorpha  canescens         .          .          .          .          .          .  192 

Kuhnistera  occidentalis  ( ?)                   .          .          .  .26 

Solidago  rupestris            .          .          .          .          .          .  1 8 

Artemisia  gnaphalodes       .          .          .          .          .  .17 

Solidago  mollis      .          .          .          .          .          .          .  12 

Solidago  rigida  .......         3 

Helianthus  rigidus          ......  2 

The  following  count,  made  in  the  buffalo  grass  formation, 
about  two  miles  beyond  the  one  last  set  forth,  shows  the  latter 
formation  as  affected  by  the  near  proximity  of  the  prairie  gras, 
formation.  The  number  of  secondary  species,  small  as  it  is,  is 
very  large  for  that  formation.  Where  this  count  was  mades 
the  dominant  grass  was  Bouteloua  oltgostachya.  The  only 
other  grass  was  Schedonnardus  -paniculatus ,  represented  by  two 
small  patches. 

Plantago  Purshii    .......  42 

Solidago  mollis   .          .          .          .          .          .          .  23 

Lygodesmia  juncea           ....'..  5 

Lepachys  columnaris  (dwarf)       .....  3 

Eriocarpum  spinulosum  ......  2 

Grindelia  squarrosa   .          .                     .                     .  .        I 

Kuhnia  glutinosa    .......  i 

Lesquerella  argentea             .          .          .          .          .  .        i 

The  constant  diminution  in  the  number  and  abundance  of 
secondary  species  as  one  passes  from  the  prairie  grass  formation 
of  the  prairie  region  to  the  buffalo  grass  and  bunch  grass  forma- 
tions of  the  transition  area  and  of  the  sand  hill  region  is  well 
illustrated  by  these  figures.  The  difference  betweeen  the 
prairie  grass  formation  in  its  ordinary  situations  and  in  the 
transition  area,  and  between  the  buffalo  grass  formation  of  the 
"range"  and  the  same  formation  in  the  transition  area  is  better 

O 

shown  by  figures  obtained  from  such  enumerations  than  in  any 
other  way.  Many  other  examples  of  the  efficacy  of  this  method 
in  representing  changes  in  the  floral  covering  as  one  passes 
from  one  district  to  another  might  be  given. 

The  method  of  actual  enumeration  of  the  individual  plants 
present  in  plots  of  a  given  size  makes  accurate  limitation  of  the 
several  grades  of  abundance  possible.  Of  course,  this  has  noth- 


Pound  and  Clements:    METHOD  OF  DETERMINING  SPECIES.    23 

ing  to  do  with  the  mode  of  disposition  of  individuals.  But  given 
a  copious,  gregario-copious  or  sparse  species,  there  still  remains 
something  more  to  be  said  before  the  abundance  of  the  species 
is  fairly  indicated.  Collation  of  the  results  of  a  large  number 
of  enumerations  has  shown  that  six  grades  of  copious  plants 
may  be  recognized  readily.  The  first,  in  which  the  average 
number  of  individuals  in  a  plot  five  meters  square  exceeds  200, 
corresponds  to  copious.1  As  examples,  there  may  be  cited  from 
the  prarie  formations  Amorpha  canescens,  with  an  average  of 
309  in  the  prairie  region,  Aster  multiflorus  with  an  average  of 
275  in  the  prairie  region  and  about  230  in  the  sand  hill  region ; 
from  the  herbaceous  layer  of  woody  formations,  Verbesina 
alternifolia  (which  is  almost  gregarious  at  times),  with  an  aver- 
age of  245.  To  the  second  degree  (copious2)  those  species  may 
be  assigned  in  which  the  average  number  of  individuals  in  a 
plot  is  from  150  to  200,  such  as  Plantago  Purshii  (162)  in  the 
Peppergrass-Cactus  formation  in  the  transition  area  between 
the  sand  hill  region  and  the  foot  hill  region.  Those  species 
with  an  average  ranging  from  TOO  to  150  may  be  assigned  to 
the  third  degree  (copious3).  Examples  are  :  Aster  sagittifolius , 
which  has  an  average  of  133  in  the  herbaceous  layer  of  the  Bur- 
oak-Elm- Walnut  formation  in  the  Mississippi  basin  region  and 
Solidago  rupestriS)  which  has  an  average  of  104  in  the  Sporo- 
bolus-Koeleria-Panicum  formation  in  the  prairie  region.  In 
the  fourth  degree  (copious4)  those  species  may  be  included 
which  have  an  average  of  from  50  to  100,  such  as  Glycorhiza 
lepidota  in  the  river  valleys  in  the  sand  hill  region,  where  its 
average  is  83.  All  of  the  foregoing  are  of  sufficient  abundance 
to  be  included  in  the  general  term  "  copious,"  taking  the  latter 
to  represent  a  quantitative  idea  as  well  as  the  manner  of  associ- 
ation of  the  individuals.  Where  the  average  falls  below  50  and 
exceeds  5,  we  call  the  species  "  subcopious."  Comparison  and 
collation  of  statistics  has  shown  that  subcopious  species  fall  into 
two  groups,  in  one,  which  we  call  subcopious,1  the  average 
does  not  fall  below  15.  Examples  are  :  Kuhnistera  Candida  in 
the  Sporobolus-Koeleria-Panictim  formation  in  the  prairie  re- 
gion, where  it  has  an  average  of  18,  Solidago  mollis  in  the 
Peppergrass-Cactus  formation  in  the  transition  area  between 
the  sand  hill  and  foot  hill  regions,  where  its  average  is  slightly 
over  20,  and  Artemisia  gnaphalodes  in  the  transition  between 
the  prairie  and  the  sand  hill  regions,  where  its  average  is  16. 


24  MINNESOTA    BOTANICAL    STUDIES. 

Where  the  average  number  in  a  plot  is  between  5  and  15,  the 
species  is  called  subcopious.2  A  glance  at  the  list  given  above 
will  show  that  these  are  often  very  striking  components  of  the 
prairie  formations.  Finally,  in  case  the  average  is  below  5  and 
above  .01,  or  one  individual  in  ten  plots,  the  species  is  called 
"sparse."  Gregario-copious  species  may  be  treated  in  the 
same  way,  giving  gregario-copious,1  etc.  Antennaria  campes- 
tris  in  the  prairie  grass  formations  of  the  prairie  region,  aver- 
aging 12  patches  and  145  individuals  per  plot,  would  be  gre- 
gario-copious.3 

Although  this  method  involves  no  little  labor,  especially  when 
applied  to  social  species,  as  we  have  been  able  to  do  success- 
fully in  some  cases,  such  as  the  Peppergrass  in  the  Pepper- 
grass-Cactus  formation,  it  has  furnished  results  which  amply 
reward  the  time  and  work  required.  By  means  of  such  enum- 
erations we  have  been  able  to  determine  many  questions  with 
certainty  which  could  only  be  guessed  at  otherwise,  and  we 
have  been  able  to  make  more  accurate  limitations  of  the  regions 
and  particularly  the  transition  areas  than  we  had  thought  pos- 
sible. 


III.     LIST  OF  FRESH-WATER  ALGAE  COLLECTED 
IN   MINNESOTA   DURING    1896   AND    1897. 


JOSEPHINE  E.  TILDEN. 


During  the  past  two  years  no  special  effort  has  been  made  to 
collect  the  algae  of  the  State.  Several  species  have  been  given 
particular  study  in  the  laboratory  and  a  few  others  have  inci- 
dentally been  brought  to  notice.  The  list  comprises  only  those 
not  heretofore  recorded  in  Minnesota,  and  is  a  continuation  of 
the  series  begun  in  Vol.  I.  of  this  publication.  Attention  may 
be  called  to  the  comparatively  large  number  of  lime-secreting 
forms. 

HELMINTHOCLADIACEJE    (HARV.)   SCHMITZ   Syst.  Uebers. 
Florid,  in  Flora  4.      1889. 

240.  Chantransia    pygmaea  (KG.)  SIRDT.      Les  Batracho- 

spermes.     244,  245.      1884. 

Together  with  Chaetophora  calcarea,  Dicothrix  cal- 
carea, Lyngbya  martensiana  var.  calcarea  and  JL. 
nana,  forming  the  calcareous  crust  on  sides  of  old 
tank  and  on  twigs  in  the  water. 

Minneapolis,  Minnesota.     October  i,  1895. 

241.  Chantransia  expansa  WOOD.     Contr.  Hist.  Fresh- Water 

Algae  North  Am.  215.  pi.  19.  f.  2.     1872. 
On  stones  under  waterfall.     Osceola,  Wisconsin.     Sep- 
tember 15,  1897. 

CHARACEAE  RICHARD  in  Humb.  et  Bonpl.  Nov.  G.  I.     1815. 

242.  Chara  contraria  A.  BR.  Schweizer.     Char.  15.     1847. 
In  ditches.     Osceola,  Wisconsin.     August  31,  1895. 

243.  Chara  foetida  A.  Br.  Ann.  Sci.  Nat.  Bot.  II.  i:  354. 

1834. 

In  pool  formed  by  spring  water.  Osceola,  Wisconsin. 
September  15,  1897. 


26  MINNESOTA    BOTANICAL    STUDIES. 

ULOTRICHIACEAE  (Kc.)  BORZI  em.    DE  TONI.  Syll.  Alg.  i : 
151.     1889. 

244.  Hormiscia   zonata   (WEB.    and    MOHR)   ARESCH.   var. 

valida  (NAG.)   RABENH.    Fl.    Eur.  Algar   3:    362. 
1868. 

On  rocks  wet  with  surf.  Grand  Marais,  Lake  Su- 
perior, Minnesota.  Coll.  A.  H.  Elftman.  July  27, 
1896. 

PALMELLACEAE   (DECNE.)   NAG.  em.  DE  TONI.  Syll.  Alg. 
i:    559.      1889. 

245.  Scenedesmus  obliquus  (TURP.)  KG.  Syn.  Diat.  in  Lin- 

naea.  8  :  609.     1833. 

Grown  in  aquarium  in  which  water  was  saturated  with 
nitrous  oxide.  University  of  Minnesota,  Minneapolis, 
Minnesota.  Emil  Sandsten.  February  23,  1898. 

246.  Chlorochytrium  archerianum  HIERON.  in  Jahres.  Schles. 

Gesellsch.  296.     1887. 

In  cells  of  Sphagnum  which  had  been  kept  in  the  Uni- 
versity plant-house  six  weeks.  Osceola,  Wisconsin. 
Coll.  Conway  MacMillan.  September  15,  1896. 

247.  Tetraspora  cylindrica  (WAHLENB.)  AG.  Syst.  Alg.  188. 

no.  2.      1824. 

Attached  to  lake  bottom,  abundant  around  the  outside 
harbor  rocks.  Grand  Marais,  Lake  Superior,  Minne- 
sota. Coll.  A.  H.  Elftman.  July  27,  1896. 

248.  Palmella  miniata  LEIBL.  var.  aequalis  NAG.  Einzell. 

67.  pi.  4.  D.  f.  2.     1849. 
On    submerged    rocks    and    pebbles    in    slow   current. 

Minnehaha    creek,    Soldiers'    Home,    Minneapolis, 

Minnesota.     September  27,  1896. 
This  species  contains  calcium  carbonate  in  quantity.     It 

is  accompanied   by  filaments  which  much    resemble 

Stigeoclonium  in  its  transition  stage. 

ZYGNEMACEAE  (MENEGH.)  RABENH.  Fl.  Eur.  Algar.  2  :  228. 
1868. 

249.  MougeotiaparvulaHASs.  var.  angusta(HAss.)  KIRCHN. 

Alg.  Schles.  128.     1878. 


Tilden:   LIST  OF  FRESH-WATEK  ALGAE.  27 

Grown  in  aquarium  in  which  water  was  saturated  with 
nitrous  oxide.  University  of  Minnesota,  Minneapolis, 
Minnesota.  Emil  Sandsten.  February  23,  1898. 

RIVULARIACEAE  RABENHORST  Fl.  Eur.  Algar.  2:  2.     1865. 

250.  Calothrix  parietina  (NAG.)  THUR.  Ess.  Class.  Nostoch. 

in  Ann.  Sci.  Nat.  Bot.  VI.  I  :  381.     1875. 
On  stone  sides  of  fountain,  breaking  up  in  small  frag- 
ments when    peeled    off.     Kenwood,    Minneapolis, 
Minnesota.     August  3,  1895. 

251.  Dichothrix  calcarea  TILDEN  Am.  Alg.  Cent.  II.  no.  165. 

1896.     Bot.  Gaz.  23 :  95-104.     pi.  7-9.     F.   1897 
Forming  a  part  of  the  lime  incrustation  which  covers 
sides  of  wooden  tank.     With  no.  240.     Minneapolis, 
Minnesota.     October  i,  1895. 

252.  Rivularia   biasolettiana   MENEGH.    in   Zanardini  Syn. 

Alg.  in  mari  Adriatico  collect,  in  Reale  Acad.  Sci. 
Torino.  II.  IV.  42.      1841. 

On  rocks  at  edge  of  lake.  Big  Stone  lake,  Dakota. 
Coll.  David  Griffiths.  October  4,  1895. 

253.  Gloeotrichia   pisum   (Ac.)  THURET.     Essai  de  class. 

Nostochinees   in    Ann.  Sci.  Nat.  Bot.  VI.   i  :  382. 

,  l875- 

Floating  on  surface  of  water  in  large  quantity.  Lake 
Minnewaska,  Glenwood,  Minnesota.  Coll.  Elizabeth 
H.  Foss.  August,  1897. 

NOSTOCEAE  KUTZ  Phyc.  gen.  203.     1843. 

254.  Anabaena   azollae   STRASB.  Bot.    Prakt.    341.   f.   130. 

1887. 

In  chambers  in  the  leaves  of  Azolla  Caroliniana.  Uni- 
versity plant-house,  Minneapolis,  Minnesota.  Sep- 
tember 15,  1896. 

255.  Anabaena  cycadearum   REINKE,   Bot.   Zeit.   37:  473- 

476.  pi.  6.  f.  1-5.     1879. 

In  roots  of  Cycas  revoluta.  University  plant-house, 
Minneapolis,  Minnesota.  December  20,  1896. 

256.  Anabaena  flos-aquae  (LYNGB.)  BREB.  in  Brebissonet 

Godey.     Algues  des  environs  de  Falaise.  36.     1835. 
Floating  in   abundance  on   surface   of   water.     Cedar 


28  MINNESOTA    BOTANICAL    STUDIES. 

lake,  Hennepin  county,  Minnesota.     Coll.  Miss  M. 
G.  Fanning  and  H.  B.  Humphrey.   October  28,  1897. 

VAGINARIEAE  GOMONT    in    Morot,   Journ.   de  Bot.  4:351. 
1890. 

257.  Schizothrix  rupicola  TILDEN.  Am.  Alg.  Cent.  II.  no. 

175.    1896.    Bot.  Gaz.  23:  95-104.  pi.  7-9.  F.  1897. 
Bare  and  dry  sandstone  cliffs.     Soldiers'  Home,  Min- 
nehaha  Falls,  Minnesota.     Coll.  C.  W.  Hall.     Sep- 
tember 28,  1896. 

258.  Schizothrix  lardacea  (CESATI)  GOMONT.  Monogr.  des 

Oscill.  in  Ann.  Sci.  Nat.  Bot.  VII.  15:  311.  pi.  8.  f. 

8,  9.     1892. 

In  a  large  bottle  of  distilled  water  left  standing  for 
several  months.  Botanical  laboratory,  University  of 
Minnesota,  Minneapolis,  Minnesota.  1896.  Det. 
by  Gomont. 

LYNGBYEAE  GOMONT  Ess.  class  des  Nostocacees  homocystees 
in  Morot  Journ.  de  Bot.  4:  353.     1890. 

259.  Lyngbya  martensiana  MENEGH.  var.  calcarea  TILDEN. 

Am.    Alg.    Cent.    II.    no.    178.     1896.     Bot.    Gaz. 
23:  95-104.  pi.  7-9.  F.  1897. 

With  no.  240.  Minneapolis,  Minnesota.  October  i, 
1895. 

260.  Lyngbya  nana  TILDEN.     Am.  Alg.  Cent.  II.  no.  179. 

1896.     Bot.  Gaz.  23:  95-104.  pi.  7-9,  F.  1897. 
With  no.  240.     Minneapolis,  Minnesota.     October    i, 
1895. 

261.  Phormidium  valderianum  (DELP.)  GOMONT  Monogr. 

des  Oscill.  in  Ann.  Sci.  Nat.  Bot.  VII.  16 :  167.  pi. 

4.  f.  20.     1892. 
In   arm  of   Mississippi  river    (old  channel),  St.  Paul 

Park,  Minnesota.     Coll.   E.  M.   Freeman,    October 

3,  1897. 
With   Oscillator ia  geminata,  O.  tennis  var.  tergestina, 

Phormidium  valderianum,  and  species  of   Chroococ- 

cus,  Palmellaceae,  Coccocklorts,  J^haphtdmm,  Poly- 

cystis  and  Scenedemtis .     Det.  by  Gomont. 

262.  Oscillatoria  geminata  MENEGH.  Consp.  Algol,  euganeae. 

9.  1837. 
With  no.  261. 


Tilden:    LIST  OF  FRESH-WATER  ALGAE.  29 

263.  Oscillatoria  tenuis  AGARDH.  Alg.  Dec.  2  125.     1813. 
With  no.  261. 

264.  Gloeocapsa  calcarea  n.  sp. 

Forming  a  calcareous  crust,  light  gray  to  light  aerugi- 
nous  in  color,  2-3  mm.  in  thickness ;  cells  6-9  mic. 
in  diameter,  4-16  united  in  families ;  families  25-50 
mic.  in  diameter ;  sheath  colorless,  somewhat  thin ; 
cell-contents  aeruginous,  granular. 

Associated  with  several  other  lime-secreting  algal  forms. 

On  boards  where  spring  water  from  trough  drips  down 
constantly.  Osceola,  Wisconsin.  September  15,  1897. 


IV.   CORRECTIONS    AND  ADDITIONS  TO  THE 
FLORA  OF  MINNESOTA. 


A.  A.  HELLER. 


Chenopodium  Boscianum  MOQ^  Enum.  Chenop.  21.    1840. 

This  species  should  be  stricken  from  the  list  of  Minnesota 
plants,  as  no  specimen  of  it  has  yet  been  reported  from  the 
State.  Sheldon's  "  1555,  Lake  Benton,"  as  well  as  specimens 
from  other  localities,  quoted  in  the  Metaspermas  of  the  Minne- 
sota Valley,  probably  belong  to  Chenopodium  album. 

Sophia  pinnata  (WALT.)  BRITTON,  111.  Fl.  2  :  145.     1897. 
Erysimum  pinnatum  WALT.  Fl.  Car.   174.      1788. 
Sisymbrium  canescens  NUTT.  Gen.  2:    68.      1818. 
Descurainia  pinnata    BRITTON,    Mem.    Torr.  Bot.  Club,  5  : 

173.      1894. 

The  plants  referred  to  this  species  in  the  Metasperma?  of  the 
Minnesota  Valley,  under  the  name  of  "  Sisymbrium  rnultifidum," 
belong  to  Sophia  incisa,  with  the  exception  of  Sheldon's  "  1406, 
Lake  Benton,"  and  "Taylor,  1044,  Glenwood,"  which  are 
specimens  of  Sophia  Hartivegiana.  Sophia  pinnata  does  not 
occur  in  Minnesota. 

Potentilla  leucocarpa  RYDBERG,  in  111.  Fl.  2  :  212.     1897. 

To  this  species  belongs  the  specimen  collected  by  Sheldon  at 
Fergus  Falls,  Otter  Tail  county,  August,  1892,  and  referred  by 
him  to  Potentilla  Nicolletii.  Another  specimen,  also  collected 
by  Sheldon,  and  determined  as  P.  Nicolletii,  was  obtained  at 
Silver  Lake,  Otter  Tail  county,  September,  1892. 

Potentilla  Monspeliensis  L.  Sp.  PI.  499.     1753. 

A  specimen  of  this  was  collected  at  Pelican  Lake,  Otter  Tail 
county,  August,  1892,  by  Sheldon,  but  referred  to  Potentilla 
Nicolletii,  in  Minn.  Bot.  Stud.  I  :  16. 


Heller:   THE  FLORA  OF  MINNESOTA.  31 

Potentilla  Nicolletii  (WATS.)  SHELDON,  Minn.  Bot.  Stud,  i : 

16.     1894. 
Potentilla  supina  var.  Nicolletii  WATS.  Proc.  Am.  Acad.  8  : 

553-      1873. 

As  shown  by  the  two  preceding  notes,  this  species  does  not 
occur  in  Minnesota,  for  the  specimens  upon  which  Mr.  Sheldon 
raised  the  variety  to  specific  rank,  belong  to  another  species,  or 
rather  to  two  species. 

Potentilla    pentandra    ENGELM.  :    T.    &    G.    Fl.    N.    A. 

i :  447.     1840. 

Not  previously  reported  from  Minnesota.  Collected  at  Jor- 
dan, Scott  county,  June,  1891,  by  C.  A.  Ballard,  no  252.  In 
the  Metaspermas  of  the  Minnesota  Valley,  this  specimen  is  re- 
ferred to  Potentilla  Canadensis,  a  species  which  it  does  not  at 
all  resemble. 

Malus  loensis  (Wooo)  BRITTON,  111.  Fl.  2:235.     1897. 

Pyrus  coronaria  var.  loensis  WOOD,  Class  Book,  333.     1860. 
Pyrus  loensis  BAILEY,  Am.  Gard.  12:473.      1891. 

In  the  Metaspermas  of  the  Minnesota  Valley  this  species  is 
included  under  "  Pirus  coronaria"  and  the  following  speci- 
mens cited  there  belong  to  it,  and  not  to  Malus  coronaria:  Bal- 
lard, 345,  Helena,  Scott  County  ;  Sheldon,  659,  Waseca  ;  Sand- 
berg,  Red  Wing.  Sheldon's  322,  Smith's  Mills,  Blue  Earth 
county,  may  belong  here,  but  the  specimen  is  so  mutilated  and 
imperfect  that  accurate  determination  from  it  alone  is  not 
possible. 

Geranium  Bicknellii   BRITTON,    Bull.  Torr.    Bot.    Club, 

24:  92.     1897. 

Apparently  common  in  Minnesota,  as  evinced  by  the  follow- 
ing collections :  J.  H.  Sandberg,  Taylor's  Falls,  Two  Harbors, 
Red  Wing ;  C.  L.  Herrick,  Minneapolis,  St.  Louis  river ;  F. 
F.  Wood,  Pike  Lake ;  L.  H.  Bailey,  Vermilion  Lake,  no. 
199;  T.  S.  Roberts,  Duluth ;  C.  A.  Ballard,  Prior's  Lake, 
Cleary's  Lake,  Scott  county ;  B.  C.  Taylor,  Alexandria,  Tay- 
lor's Falls;  Otto  Lugger,  Tower;  E.  P.  Sheldon,  Milaca, 
Mille  Lac  Reservation,  Nichols.  It  has  heretofore  been  con- 
fused with  Geranium  Carolinianum,  but  that  species  does  not 
seem  to  have  been  collected  in  Minnesota,  as  no  specimens 
from  the  State  are  found  in  the  herbarium  of  the  University. 


32  MINNESOTA    BOTANICAL    STUDIES. 

Lechea  stricta  LEGGETT  ;  Britton,  Bull.  Torr.  Bot.  Club, 
21  :    251.      1894. 

The  Illustrated  Flora  gives  the  range  of  this  species  as 
"Wisconsin,  Illinois,  Iowa."  It  has  also  been  collected  at 
several  localities  in  Minnesota.  We  have  two  specimens  col- 
lected by  J.  H.  Sandberg,  one  at  Sandy  Lake,  August,  1891, 
labeled  "  Lechea  minor"  the  other  at  Centreville,  July  30, 
1891,  labeled  "  Lechea  Leggettii"  There  is  also  a  specimen 
collected  at  Zumbrota,  August,  1892,  by  C.  A.  Ballard,  labelled 
"  Lechea  Leggettii"  and  one  collected  by  Miss  Jennie  E.  Camp- 
bell, at  Rockville,  July,  1896.  E.  P.  Sheldon  also  collected  it 
at  St.  Croix  Falls,  Wisconsin,  September,  1892. 

The  type  of  this  species  was  presumably  collected  by  Mr. 
M.  S.  Bebb,  as  we  have  a  specimen  from  his  collection,  with 
the  record :  "  Lechea  stricta,  Leggett  ms.  .  .  .  Fountaindale, 
Winnebago  County,  Illinois,  1879."  By  referring  to  the  Bo- 
tanical Gazette,  15  :  308.  1890,  I  find  that  the  species  was  re- 
ported from  Minnesota,  previous  to  its  publication,  by  E.  J. 
Hill,  who  says:  "The  only  Lechea  seen  was  one  called  by 
Mr.  W.  H.  Leggett,  who  has  given  special  attention  to  these 
plants,  L.  minor  Lam.  var  stricta.  It  grows  on  sandy  hills, 
especially  those  thinly  covered  with  Pinus  Banksiana  and  P. 
resinosa,  where  the  ground  is  not  too  much  shaded,  and  on 
rocky  hills  and  ledges  with  a  thin  covering  of  soil,  on  top  of 
which,  Jasper  Peak,  the  highest  point  in  the  vicinity  of  Tower, 
I  find  it  common." 


V.     NEW   AND    INTERESTING    SPECIES    FROM 
NEW  MEXICO. 


A.  A.  HELLER. 


Allionia  diffusa  n.  sp. 

Stems  terete,  two  or  three  from  a  perennial  rootstalk,  diffusely 
branched  from  the  base,  20  to  30  cm.  in  length,  whitened  and 
glabrous  below,  the  middle  part  usually  marked  with  several 
lines  of  short,  curved  hairs,  the  branches  immediately  below  the 
inflorescence,  as  well  as  the  inflorescence  itself,  covered  with 
spreading,  glandular  hairs ;  leaves  sessile,  lanceolate-linear, 
slightly  narrowed  at  the  base,  the  lowest  5  to  6  cm.  long,  the 
upper  ones  about  half  that  length,  all  acute,  the  upper  face  pro- 
vided with  a  grayish  margin,  midvein  prominent ;  involucres 
clustered  at  the  ends  of  the  branches,  mature  ones  about  7  mm. 
across,  their  lobes  triangular-lanceolate,  acutish,  between  2  and 
3  mm.  long;  perianth  pale  rose  color,  6  mm.  long,  its  lobes 
broadly  obovate ;  stamens  three,  these,  as  well  as  the  style,  ex- 
serted. 

The  type  is  our  no.  3740,  collected  June  21,  1897,  on  dry, 
gravelly  hills,  ten  miles  west  of  Santa  Fe,  altitude  6000  feet. 
The  diffuse  habit  of  the  plant  cannot  always  be  well  shown  in 
dried  specimens,  but  it  is  quite  marked  in  the  living  state,  the 
lower  branches  being  almost  procumbent.  This  species  is,  per- 
haps, plentiful  in  the  vicinity  of  the  place  where  it  was  first 
found,  but  as  only  one  trip  was  made  to  that  place  after  it  came 
into  bloom,  only  a  few  specimens  were  collected. 

Pedicularis  fluviatilis  n.  sp. 

Stems  several  from  a  perennial  root,  erect,  15  to  20  cm.  high, 
lanate  pubescent,  especially  above,  leafy,  or  the  upper  part 
somewhat  naked;  leaves  alternate,  rather  distant,  dark  green, 
mature  ones  5  to  6  cm.  long,  i  cm.  wide,  linear-oblong  in  out- 
line, acute  or  acutish,  deeply  pinnately  parted,  the  lobes  of 


34  MINNESOTA    BOTANICAL    STUDIES. 

almost  uniform  width,  and  lobed  or  serrate  with  spreading 
teeth ;  spikes  leafy  bracteate,  dense,  4  to  6  cm.  long,  and  almost 
as  broad ;  calyx  i  cm.  long,  obliquely  cleft  to  the  base  on  the 
lower  side,  the  upper  side  notched  with  a  shallow  rounded 
sinus ;  corolla  lemon  yellow  or  faintly  purple  tinged,  slightly 
over  2  cm.  in  length,  6  mm.  wide,  summit  of  the  galea  incurved, 
the  tip  provided  with  two  cusps. 

The  type  is  our  no.  3639,  collected  June  2,  1897,  in  a  meadow 
nine  miles  east  of  Santa  Fe,  altitude  8000  feet.  The  name 
fluviatilis  is  not  very  appropriate,  but  as  specimens  have  been 
distributed  under  this  name,  I  consider  it  better  policy  to  de- 
scribe it  under  the  name  it  has  borne,  rather  than  cause  con- 
fusion by  assigning  another  more  appropriate  one.  The  speci- 
mens were  growing  in  a  grassy  meadow,  on  the  banks  of  the 
Santa  Fe  creek,  opposite  "Monument  Rock."  For  some 
months  of  the  year  this  meadow  is  overflowed  by  water  from  an 
irrigating  ditch,  so  that  then  the  plants  are  actually  growing  in 
water,  as  indicated  by  the  name.  This  species  seems  to  be 
closely  related  to  the  common  eastern  P.  Canadensis. 

Pentstemon  caudatus  n.  sp. 

Glabrous  throughout ;  stems  herbaceous,  rather  stout  and 
fleshy,  simple,  erect,  25  to  30  cm.  high,  pruinose,  leafy  through- 
out ;  leaves  all  sessile,  more  or  less  clasping,  pruinose,  rather 
fleshy,  margined  with  a  narrow  pale  line,  the  lower  ones  spat- 
ulate-oblong,  or  oblong-lanceolate,  moderately  acute,  4  to  7 
cm.  long,  5  mm.  to  r  cm.  wide,  gradually  becoming  longer  and 
broader  until  the  inflorescence  is  reached,  where  they  are 
broadly  ovate- lanceolate,  with  long  and  narrow  tips  ;  beginning 
with  the  inflorescence  they  become  gradually  shorter,  but  still 
retain  the  broad  base,  until  near  the  very  end  of  the  stalk,  where 
they  are  reduced  to  lanceolate  bracts  :  flowers  pale  violet  or 
pinkish,  numerous  in  thyrsiform  close  clusters  in  the  axes  of  the 
leaves,  occupying  from  one-half  to  three-fourths  of  the  length 
of  the  stem :  calyx  about  5  mm.  long,  its  lobes  lanceolate,  or 
ovate-lanceolate,  long-pointed,  scarious  margined,  broadly  so 
near  the  base;  corolla  slightly  over  2  cm.  in  length,  the  tube 
gradually  dilated  into  the  funnelform  throat,  the  orbicular-obo- 
vate  lobes  nearly  equal,  about  7  mm.  long,  spreading;  sterile 
filament  bearded  in  the  upper  half  on  one  side,  the  tip  some- 
what dilated  and  curled ;  anthers  dehiscent  from  base  to  apex, 
puberulous  at  line  of  dehiscence. 


Heller:    NEW  AND  INTERESTING  SPECIES  FROM  MEXICO.       35 

The  type  is  our  no.  3580,  collected  May  26,  1897,  at  Bar- 
ranca, Taos  county,  altitude  6900  feet.  It  is  very  abundant  in 
open  grassy,  sandy  soil,  about  Barranca  station,  growing  in  large 
patches.  This  species  is  likely  to  occur  in  Colorado  also,  as  it 
occurs  on  the  plateau  which  runs  up  into  Colorado  between  the 
two  mountain  ranges,  and  specimens  of  it  will  probably  be  found 
in  herbaria  under  the  name  of  P.  acuminatis  Dougl.,  which 
northwestern  species  seems  to  be  a  convenient  depository  for 
anything  which  at  all  remotely  resembles  the  original. 

Senecio  Sanguisorbae  DC.  Prodr.  6:  427.     1837. 

Specimens  referable  to  this  species  were  collected  in  Santa 
Fe  Canon,  about  twelve  miles  from  Santa  Fe.  They  were 
growing  in  wet  ground  along  the  stream,  at  a  place  well  within 
the  woods.  Although  numerous  plants  were  seen,  only  a  few 
specimens  were  obtained,  as  it  was  just  coming  into  bloom  and 
that  part  of  the  Canon  was  not  again  visited.  To  the  best  of 
my  knowledge,  the  species  has  not  hitherto  been  recorded  within 
the  borders  of  the  United  States.  No.  3820. 

Sitilias  Rothrockii  (A.  GRAY)  GREENE,  Pittonia,  2:  180.  1891. 
Pyrrhopappus   Rothrockii  A.  GRAY,  Proc.   Am.  Acad.    n  : 
So.      1876. 

In  our  distribution  of  New  Mexican  plants  of  1897,  this  spe- 
cies was  erroneously  determined  as  "Sitilias  mit Uica ulis  (DC.) 
GREENE."  It  was  collected  in  a  meadow  along  the  Rio  Grande 
river  opposite  the  Indian  Pueblo  of  San  Juan,  no.  3758.  The 
specimens  were  obtained  either  in  damp  ground,  or  sometimes 
actually  in  water  in  swampy  places.  In  some  of  the  specimens 
the  leaves  are  entire,  or  the  lower  ones  only  slightly  toothed, 
while  in  others  the  lower  ones  are  conspicuously  pinnatifid. 
The  original  came  from  "  Fisch's  Ranch,  in  southern  Arizona, 
at  5000  feet  altitude."  Rothrock,  699. 

The  type-specimens  are  deposited  in  the  herbarium  of  the 
University  of  Minnesota. 


VI.    SOME   MUSCI   OF   THE   INTERNATIONAL 
BOUNDARY. 


JOHN  M.  HOLZINGER. 


The  mosses  listed  below  were  personally  collected  during  the 
summer  of  1897  in  northeastern  Minnesota  along  the  Dawson 
canoe-route  between  Ely  and  Grand  Portage.  The  route  in- 
cludes the  following  lakes  :  Fall,  Basswood,  Newton,  Sucker, 
Carp,  Melon,  Seed,  Knife,  Otter-track,  Saganaga,  Granite, 
Gunflint,  North,  South,  Rose,  Rove,  Mountain,  Moose,  North- 
Fowl,  South-Fowl  and  Superior. 

Acknowledgements  and  thanks  are  due  to  the  following  per- 
sons for  assistance  :  To  M.  Jules  Cardot  for  determination  of 
Fontinalaceae,  to  Dr.  R.  True  for  determination  of  Dicrana,  to 
Mr.  A.  J.  Grout  for  determination  and  correction  of  the  Eu- 
rhynchia,  Brachythecia  and  Pylaisiellee,  to  Mrs.  E.  G.  Britton 
for  determination  of  Orthotricha,  to  Dr.  G.  N.  Best  for  deter- 
mination and  verification  of  Leskeae,  Thuidia,  Myurellas  and 
Anomodonta  and  some  other  Hypnaceas,  and  to  Dr.  C.  Warn- 
storf  for  determination  of  the  Sphagna. 

A  further  list,  including  additional  species,  may  be  expected 
to  follow  this  at  a  later  date. 

1.  Sphagnum  acutifolium  EHRH. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
shore  of  Fall  lake,  at  Camp  i  (June  8-10,  1897). 

2.  Sphagnum  fuscum  KLINGGR. 

Near  Port  Arthur,  Canada  (June  18,  1897). 

3.  Sphagnum  girgensohnii  Russ. 

On  the  point  of  land  at  the  foot  of  Kawasatchong  falls, 
shore  of  Fall  lake  (June  8-10,  1897). 

4.  Sphagnum  medium  LIMPR. 

At  the  farther  end  of  the  portage  around  Pipestone  rapids, 
between  Newton  and  Basswood  lakes  (June  u,  1897), 


Holzinger :    MUSCI  OF  THE  INTERNATIONAL  BOUNDARY.       37 

5.  Sphagnum  recurvum  parvifolium  SONDTN. 

With  Sphagnum  medium. 

6.  Sphagnum  teres  squarrosulum  (LEsqJ  WARNST. 

At  the  farther  end  of  the  portage  around  Pipestone 
rapids,  between  Newton  and  Basswood  lakes  (June  n, 
1897). 

7.  Sphagnum  squarrosum  PERS. 

On  the  trail  between  Eve  lake  and  fall,  near  the  base  of 
Kawasatchong  falls  (June  8-10,  1897). 

8.  Sphagnum  wulfianum  GIRO. 

Same  locality  as  S.  sqnarrosum. 

9.  Rhabdoweisia  denticulata  B.  S. 

At  the  lower  end  of  Pipestone  rapids  from  Newton  lake  to 

Basswood  lake  (June  10,  u,  1897). 
On  a  small  point  of  land  at  the  base  of  the  United  States 

peninsula,  shore  of  Basswood  lake  (June  IT,  1897). 
On  Safety  island,  Lake  Saganaga  (June  16,  1897). 

10.  Cynodontium  polycarpum  B.  S. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
shore  of  Fall  lake,  near  Camp  i  (June  8-10,  1897). 

At  the  farther  end  of  Pipestone  rapids,  shore  of  Bass- 
wood  lake  near  the  portage  (June  10,  n,  1897). 

On  the  Prairie  portage,  shore  of  Basswood  lake,  near 
the  rapids  from  Sucker  lake,  at  Camp  4  (June  12,  13, 
1897). 

u.  Dicranum  palustre  LA  PYL  (D.  boryeani  DE  NOT). 

On  the  Prairie  portage,   shore  of  Basswood  lake,  near 

the  rapids  from   Sucker  lake,  at    Camp  4    (June   12, 

13,  1897). 
On  a  small  island  in  Lake  Saganaga,  called  by  our  party 

Safety  island,  close  by  South  island,  at  Camp  8  (June 

16,  1897). 
On  the  portage  from  Mountain  lake  to  Moose  lake  (June 

20,  1897). 

12.  Dicranum  palustre  alatum  BARNES. 

On  Safety  island,  in  Lake  Saganaga.  With  the  species 
(June  16,  1897). 

13.  Dicranum   drummondii    C.  MULL. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
shore  of  Fall  lake,  at  Camp  i  (June  8-10,  1897). 


38  MINNESOTA    BOTANICAL    STUDIES. 

At  the  lower  end  of  Pipestone  rapids,  on  Basswood  Lake, 

near  Camp  2    (June  10,  n,  1897). 
On   the  Prairie  portage,  shore  of  Basswood  lake,  near 

the  rapids  from   Sucker  lake,   at   Camp  4   (June  21, 

13,  1897). 
On  Safety  island,  Lake  Saganaga,  at  Camp  8   (June  16, 

1897). 
On  the  portage  from  Mountain  lake  to  Moose  lake  (June 

20,    1897). 

On  Grand  Portage  island,  north  shore  of  Lake  Superior 

(June  23,  1897). 

14.  Dicranum  flagellare  HEDW. 

On  the   way  from  Ely  to  Winton,   shore    of   Fall  lake 

(June  8,  1897). 
On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 

shore  of  Fall  lake,  at  Camp  i   (June  8-10,  1897). 
At  the  lower  end  of  Pipestone  rapids,  on  Basswood  lake, 

near  Camp  2  (June  10,  u,  1897). 
On  Safety  island,  Lake  Saganaga,  at  Camp  8  (June  16, 

1897). 
On  the  portage  from  North  lake  to  South  lake,  the  divide 

between  the  waters  of  Hudson  Bay  and  Lake  Superior 

(June  20,  1897). 
On  Grand  Portage  island,  north  shore  of  Lake  Superior 

(June  23,  1897). 

15.  Dicranum  fuscescens  TURN. 

At  the  lower  end  of  Pipestone  rapids,  on  Basswood  lake, 

near  Camp  2  (June  10,  n,  1897). 
On  the  Prairie  portage,  shore  of  Basswood  lake,  near 

the   rapids   from   Sucker  lake,   at  Camp  4  (June    12, 


At  the  south  end  of  Gunflint  lake,  at  Camp    10  (June 

20,    1897). 

On  Grand  Portage  island,  north  shore  of  Lake  Superior 

(June  23,  1897). 

16.  Dicranum  longifolium  HEDW. 

At  the  lower  end  of  Pipestone  rapids,  on  Basswood  lake, 

near  Camp  2  (June  10,  n,  1897). 

On    a    small  point  of   land    at   the  base  of   the  United 
States  peninsula,  Basswood  lake  (June  n,  1897). 


Holzinger:    MUSCI  OF  THE  INTERNATIONAL  BOUNDARY.        39 

On  Basswood  lake,   at  the  farther  end  of   the  portage 

across  the  United  States  peninsula  (June  12,  1897). 
On  the  Prairie  portage,  shore  of    Basswood  lake,  near 

the  rapids  from  Sucker  lake,   at  Camp  4  (June    12, 

13,  1897). 
On  Safety  island,  Lake  Saganaga,  at  Camp  8  (June  16, 

1897). 
On  the  portage  from  North  lake  to  South  lake  (June 

20,  1897). 

On  the  portage  from  Mountain  lake  to  Moose  lake  (June 

21,  1897). 

Dicranum  montanum  HEDW. 

On  the  point  of  land  at  the  base  of  Kawastachong  falls, 

shore  of  Fall  lake,  at  Camp  i  (June  8,  9,  10,  1897). 
At  the  lower  end  of  Pipestone  rapids,  on  Basswood  lake, 

near  Camp  2  (June  10,  u,  1897). 
At   the    farther  end    of   the    portage  across  the  United 

States  peninsula,  on  Basswood  lake  (June  12,  1897). 
On  the  Prairie  portage,  shore  of   Basswood  lake,  near 

the  rapids  from  Sucker  lake,   at  Camp  4  (June    12, 

i3»  1897). 

On  Safety  island,  Lake  Saganaga  (June  16,  1897). 
At  the  east  end  of  Gunflint  lake  (June  20,  1897). 

Dicranum  scoparium  HEDW. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
on  the  shore  of  Fall  lake  (June  8-10,  1897). 

Dicranum  undulatum  EHRB. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 

shore  of  Fall  lake  (June  8-10,  1897). 
On  the  Prairie  portage,  shore  of  Basswood  lake,  near 

the  rapids  from   Sucker  lake,  at  Camp  4  (June  12, 

i3»  1897). 
On  the  east  end  of  Gunflint  lake,  at  Camp  10  (June  20, 

1897). 
On  the  portage  from  North  lake  to  South  lake,  the  divide 

between  Hudson  Bay  and  Lake  Superior  (June    20, 

1897). 

Dicranum  viride  B.  S. 

Along  the  road  from  Ely  to  Fall  lake  (June  8,  1897). 
On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
shore  of  Fall  lake,  at  Camp  i  (June  8-10,  1897). 


40  MINNESOTA    BOTANICAL    STUDIES. 

On  the  Prairie  portage,  shore  of  Basswood  lake,  near 
the  rapids  from  Sucker  lake,  Camp  4  (June  12,  13, 
1897). 

21.  Fissidens  incurvus  SCHW. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
shore  of  Fall  lake,  near  Camp  i  (June  8-10,  1897). 

22.  Fissidens  osmundoides  HEDW. 

Locality  same  as  last. 

23.  Leucobryum  glaucum  SCH. 

Near  Camp  2,  at  the  lower  end  of  Pipestone  rapids,  on 
Basswood  lake  (June  10,  n,  1897). 

24.  Ceratodon  purpureus  BRID. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
shore  of  Fall  lake,  near  Camp  i  (June  8-10,  1897). 

At  the  farther  end  of  the  portage  around  Pipestone  rap- 
ids, shore  of  Basswood  lake  (June  10,  u,  1897). 

Along  the  portage  from  Mountain  lake  to  Moose  lake 
(June  20,  1897). 

25.  Distichium  capillaceum  B.  S. 

At  the  base  of  Kawasatchong  falls,  shore  of  Fall  lake, 

Camp  i  (June  8-10,   1897). 
At  the  base  of   the  United  States  peninsula,  basswood 

lake  (June  n,  1897). 
On  Grand  Portage  island,  north  shore  of   Lake  Superior 

(June  23,  1897). 

26.  Barbula  ruralis  HEDW. 

At  the  farther  end  of  the  portage  across  the  United  States 
peninsula,  shore  of  Basswood  lake  (June  12,  1897). 

27.  Barbula  tortuosa  W.  and  M. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
shore  of  Fall  lake  near  Camp  i  (June  8-10,  1897). 

On  Grand  Portage  island,  north  shore  Lake  Superior 
(June  23,  1897). 

28.  Grimmia  apocarpa  HEDW. 

On  the  Prairie  portage,  shore  of  Basswood  lake,  near  the 
rapids  from  Sucker  lake,  Camp  4  (June  12,  13,  1897). 

29.  Hedwigia  ciliata  EHRH. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
shore  of  Fall  lake,  near  Camp  i  (June  S-io,  1897). 


Holzinger :   MUSCI  OK  THE  INTERNATIONAL  BOUNDARY.        41 

On  a  point  of  land,  at  the  base  of  the  United  States  pe- 
ninsula, shore  of  Basswood  lake  (June  n,  1897). 

On  Safety  island,  Lake  Saganaga  (June  16,  1897). 

On  the  portage  from  South-Fowl  lake  to  Pigeon  river 
(June  21,  1897). 

30.  Amphoridium  lapponicum  SCH. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
shore  of  Fall  lake,  near  Camp  i  (June  8-10,  1897). 

31.  Ulota  crispa  BRID. 

In  the  woods  along  the  road  from  Ely  to  Winton,  on  Fall 
lake  (June  8,  1897). 

32.  Ulota  curvifolia  BRID. 

On  a  small  point  of  land  at  the  base  of  the  United  States 
peninsula  (June  n,  1897). 

33.  Ulota  hutchinsiae  Sen. 

On  Safety  island,  in  Lake  Saganaga  (June  16,  1897). 

34.  Orthotrichum  speciosum  NEKS. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
shore  of  Fall  lake,  near  camp  i  (June  8-10,  1897). 

NOTE  :  This  plant  agrees  in  appearance  with  plants 
from  Idaho  and  Washington,  except  that  the  leaves  are 
only  slightly  papillose,  the  papillae  being  mostly  low  and 
simple,  exactly  as  figured  for  O.  elegans  Schwaegr.,  in 
Husnot,  Muse.  Gall.  Another  point  of  departure  is  the 
smooth  or  nearly  smooth  capsule.  In  these  two  points 
it  seems  to  approach  O.  elegans.  Yet  the  disposition  of 
the  cilia  of  the  peristome  is  not  as  described  in  this  spe- 
cies, but  as  in  O.  speciosum.  The  plant  seems  therefore 
to  stand  intermediate  between  O.  'speciosum  and  O.  ele- 
gans. And  in  that  case  Schwagrichen's  species  is  rather 
Orthotrichum  speciosum  elegans. 

35.  Orthotrichum  speciosum  roellii  VENT. 

On  trees  along  the  road  from  Ely  to  Winton,  Fall  lake. 
(June  8,  1897). 

36.  Encalypta  ciliata  HEDW. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
shore  of  Fall  lake,  near  Camp  i  (June  S-io,  1897). 


42  MINNESOTA    BOTANICAL    STUDIES. 

37.  Teraphis  pellucida  HEDW. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 

shore  of  Fall  lake,  near  Camp  i   (June  8-10,  1897). 
On  a  small  point  of  land  at  the  base  of  United  States  pe- 

insula,  shore  of  Basswood  lake  (June  u,  1897). 
At  the  lower  end  of  the  portage  around  Pipestone  rapids, 

shore  of  Basswood  lake  (June  10,  n,  1897). 
On  Safety  island,  Lake  Saganaga  (June  16,  1897). 
On  the  portage  from  South  lake  to  Rat  lake  (June  20, 

1897). 

38.  Funaria  hygrometrica  HEDW. 

On  the  portage  from  North  lake  to  South  lake,  the  di- 
vide between  Hudson  Bay  and   Lake  Superior  (June 

20,  1897). 

39.  Bartramia  oederi  SCHW. 

Along  the  road  from  Ely  to  Winton,  shore  of    Fall  lake 

(June  8,  1897). 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
shore  of  Fall  lake,  near  Camp  i  (June  8-10,  1897). 

40.  Bartramia  pomiformis  HEDW. 

On  the  lower  end  of  the  portage  around  Pipestone  rapids, 

shore  of  Basswood  lake,  near  Camp  2  (June  10,  n, 

1897). 
On  a  small  point  of  land,  at  the  base  of  the  United  States 

peninsula,  Basswood  lake  (June  n,  1897). 
On  the  Prairie  portage,  shore  of  Basswood  lake,  near 

the  rapids  from  Sucker  lake  (June  12,  13,  1897). 
On  Safety  island,  Lake  Saganaga  (June  16,  1897). 
Near  Gunflint  station  (June,  1897). 
On  the  portage  from  Mountain  lake  to  Moose  lake  (June 

21,  1897). 

On  Grand  Portage  island,  north  shore  of   Lake  Superior 
(June  23,  1897). 

41.  Leptobryum  pyriforme  Sen. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
shore  of  Fall  lake,  near  Camp  i  (June  8-10,  1897). 

42.  Webera  nutans  HEDW. 

Same  station  as  the  last. 


Hohinger :    MUSCI  OF  THE  INTERNATIONAL  BOUNDARY.       43 

43.  Mnium  cuspidatum  HKD\V. 

Same  station  as  the  last. 

44.  Mnium  punctatum  HEDW. 

On  the  road  from  Ely  to  Fall  lake  (June  8,  1897). 
At  Camp  i,  Fall  lake  (June  8-10,  1897). 

45.  Mnium  serratum  BRID. 

On  a  small  point  of  land  at  the  base  of  the  United  States 

peninsula,  shore  of  Basswood  lake  (June  n,  1897). 
On  Safety  island,  Lake  Saganaga  (June  16,  1897). 

46.  Timmia  bavarica  HESSL.  var  cucullata  (Micnx.). 

On  Grand  Portage  island,  north  shore  of  Lake  Superior 
(June  23,  1897). 

47.  Atrichum  undulatum  P.  B. 

On  the  point  of  land  at  the  base  of  Kawasatchong  falls, 
shore  of  Fall  lake,  near  Camp  i  (June  8-10,  1897). 

48.  Pogonatum  alpinum  ROELL. 

On  Grand  Portage  island,  north  shore  of  Lake  Superior 

(June  23,  1897). 

49.  Polytrichum  commune  L. 

Same  station  as  the  last. 

50.  Polytrichum  juniperinum  WILLD. 

On  the  point  of  land  at  the  base  of  Kawasatchong  lake, 
shore  of  Fall  lake,  near  Camp  i  (June  8-10,  1897). 

51.  Polytrichum  piliferum  SCHREB. 

On  the  prairie  portage,  shore  of  Basswood  lake,  near  the 
rapids  from  Sucker  lake  (June  12,  13,  1897). 

52.  Fontinalis  antipyretica  LINN. 

In  the  river  crossing  the  Grand  portage  about  four  miles 
north  of  Grand  Portage  village.     Abundant  (June  21, 

1897). 

53.  Fontinalis  duriaei  SCH. 

On  submerged  rocks  at  the  base  of  Kawasatchong  falls 
near  Camp  i  (June  8-10,  1897). 

54.  Fontinalis  holzingeri  CARDOT.  sp.  nova  in  litt. 

At   the  second  falls   of    Granite    river  ascending    from 

Lake  Saganaga  (June  17,  1897). 
"  Du   groupe    Heterophyllas,   voisine   du  F.   missourica 

Card.,  sed  foliis  rigidioribus,  red  firmo,  cellulis   longi- 


44  MINNESOTA    BOTANICAL    STUDIES. 

oribus,  haud  vel  vix  flexuosis,  valde  chlorophyllosis, 
parietibus  firmis,  distincta." 

55.  Fontinalis  hypnoides  HARTM.  "forma  foliis   apice   saepe 

denticulata." 

In  the  stream  flowing  from  North  lake  into  Little  Gun- 
flint  lake.  Abundant  at  the  lower  end  of  the  stream 
(June  20,  1897). 

56.  Dichelyma  pallescens  B.  S. 

At  the  base  of  alder  trunks  growing  along  the  bank  of 
Fall  lake,  near  Camp  i  (June  8-10,  1897). 

57.  Neckera  oligocarpa  B.  S. 

Near  Camp  i  at  the  base  of  Kawasatchong  falls,  shore 

of  Fall  lake  (June  8-10,  1897). 
On  a  small  point  of  land   near  the   base  of  the   United 

States  peninsula,  Basswood  lake  (June  n,  1897). 
At  the  farther  end  of  the  portage  across  the  United 

States  peninsula,  Basswood  lake  (June  12,  1897). 
On  Safety  island,  Lake  Saganaga  (June  16,  1897). 
On  Grand  Portage  island,  north  shore  of  Lake  Superior 

(June  23,  1897). 

58.  Neckera  pennata  HKDW. 

On  trees  along  the  road  from  Ely  to  Winton,  on  Fall 

lake  (June  8,  1897). 

Near  Camp  i,  on  Fall  lake  (June  10-12,  1897). 
On  the  Prairie  Portage,  shore  of  Basswood  lake,  near 

the  rapids  from  Sucker  lake  (June  12,  13,  1897). 

59.  Homalia  trichomanoides  jamesii  (SCHIMP.). 

Near  Camp  i,  at  the  base  of  Kawasatchong  falls,  shore 

of  Fall  lake  (June  8-10,  1897). 
On  the  portage  from  Fall  lake  to  Newton  lake  (June  10, 

n,  1897). 
On  a  small  point  of  land  at  the  base  of  the  United  States 

peninsula,  Basswood  lake  (June  n,  1897). 

NOTE  :  This  plant  has  leaves  varying  strongly  toward 
the  typical  European  form  of  the  species. 
On  Grand  Portage  island,  north  shore  of  Lake  Superior 

(June  23,  1897). 

NOTE  :  I  have  carefully  studied  the  plants  collected ; 
have  compared  them  with  Professor  Macoun's  Canadian 
specimens  sent  out  under  No.  242  ;  also  with  the  Ho- 


Holzingcr :    MUSCI  OF  THE  INTERNATIONAL  BOUNDARY.       45 

malia  collected  by  Professor  James,  near  Franconia,  N. 
H.,  named  H.  jamcsii  Schimp.,  and  probably  typical 
material  of  Schimper's  species,  also  with  H.  trichoman- 
oldcs  from  Denmark,  collected  by  Dr.  J.  Hensen,  near 
Hvako,  in  1883  ;  and  I  fail  to  find  a  single  good  reason 
for  separating  our  American  forms  of  Homalia  with  ser- 
rate leaves  from  the  European  Homalia  trichomanoides 
as  a  distinct  species.  Not  a  single  constant  character 
can  be  established  for  our  plant.  I  have  made  microm- 
eter measurements  of  the  leaf  cells  of  all  the  specimens 
examined,  and  I  find  them  essentially  of  the  same  size 
in  the  same  part  of  the  leaf  in  all  specimens.  The 
American  specimens  that  are  fertile  show  not  the  slight- 
est difference  from  European  specimens  either  in  peri- 
chastial  leaves  or  in  perforation  of  the  segments  of  the 
peristome  along  the  keel.  The  only  point  of  difference 
is  the,  on  the  whole,  more  obtuse  apex  of  the  leaves  in 
our  American  forms.  But  it  cannot  fail  to  attract  the  at- 
tention of  the  student,  especially  when  he  reviews  a  large 
number  of  forms  in  different  collections,  that  while  in 
our  American  forms  the  apex  is  on  the  whole  more 
rounded,  some  leaves  may  be  found  on  every  plant  which 
have  an  apiculate  apex.  Also  the  European  plants 
studied  show  some  leaves  more  rounded  at  the  apex  than 
others.  As  for  the  description  of  the  European  plant, 
by  European  authors  themselves,  let  me  cite  first  from 
Muscinees  de  la  France  by  M.  PAbbe  Boulay  (1884),  p. 
150.  "  Feuilles  largement  oblongue-elliptiques,  con- 
vexes-cultriformes  par  le  bord  superieur,  un  peu  repliees 
en  dessus  par  1'inflexion  du  bord  inferieur,  brevement 
apiculees,  tres  finement  denticulees  sur  tout  le  contour 
(dont  plus  grandes  et  plus  rapprochees  ver  le  sommet ; 
long.  2,  larg.  i  mm.  *****  cellules  moyennes 
8-10  fois  aussi  1.  q.  1.  ;  vers  les  bords  et  au  sommet,  elles 
sont  courtes,  rhomboidales ;  *  *  *  *  lanieres 

du  peristome  interne  lineares,  plus  longues  que  les  dents, 
peu  ou  millement  ouvertes  ver  la  carene." 

InLimpricht'sLaubmoosell  (1895), p.  715:  "Blatter 
gedrangt,  zweizerlig-abstrehend,  zuletst  abwarts  gebo- 
gen,  flach  ausgebreiten,  unsymmetrisch,  aus  herablaufen- 
der,  etwas  verschmalerter  Bariszungen-messerformung, 


46  MINNESOTA    BOTANICAL    STUDIES. 

stiimpflich,  1.8-2  mm.  lang,  und  i  mm.  breit,  am  Rande 
deroberen  Blatt  halfte  ausgefressen-gezahnt,  am  Grunde, 
an  einer  Seite  eingeschlagen  *  *  *  Innerer  Peristom 
*  *  *  in  der  linie  ritzenformig;  durch  brochen." 

c? 

Now  in  Macoun's  Cat.  of  Can.  PL,  4  (1892),  p.  163, 
the  authors  of  Homalia  macounii,  say  of  it:  "Very 
nearly  allied  to  Homalia  trichomanoides;  differs  in  the 
leaves  being  longer,  rather  lingulate,  the  lowest  basal 
cells  yellow,  the  perichaetial  leaves  more  suddenly  nar- 
rowed to  a  very  short  acumen,  the  segments  of  the  per- 
istome  cleft  between  the  articulations."  This  is  quite  all 
in  the  line  of  characterization.  Among  other  localities 
it  is  credited  to  Lake  Superior,  Drummond's  specimens 
having  been  collected  there. 

Both  from  actual  comparison  and  from  the  circum- 
stance of  locality,  the  Lake  Superior  plants  collected  by 
me  are  reasonably  referred  to  the  same  plants  upon 
which  Homalia  macounii  is  founded.  If  this  inference 
is  correct  then  the  only  valid  part  of  the  above  statements, 
which  stand  in  place  of  description,  is  the  first  phrase 
"very  nearly  allied  to  If.  trichomanoides."  The  leaf 
length  varies  according  to  European  authors  themselves. 
The  "rather  lingulate"  form  of  outline  is  ascribed  by 
Limpricht  to  Homalia  trichomanoides ,  when  he  makes  the 
leaves  "  zungen-messerformung,"  i.  e.,  "  lingulate-cultri- 
form."  As  to  the  "  lowest  basal  cells  yellow,  the  peri- 
chsetial  leaves  more  suddenly  narrowed  to  a  very  short 
acumen,"  my  own  close  observations  fail  to  verify  these 
two  characters,  which,  if  observed  by  the  authors,  must 
have  been  purely  accidental.  And  as  for  "  the  segments 
of  the  peristome  cleft  between  the  articulations,"  this 
character,  judging  both  the  European  specimens  actually 
examined,  and  from  the  painstaking  description  of  Ho- 
malia trichomanoides  made  by  European  authors  them- 
selves, as  seen  from  citations  above,  is  unconditionally 
conceded  to  belong  to  Homalia  trichomanoides  Br.  The 
only  tangible  difference,  the  slightly  more  obtuse  leaves 
it  certainly  has  in  common  with  Dr.  James'  own  speci- 
mens of  Homalia  jamesii.  If  now  we  turn  to  Lesq.  and 
James'  Manual  of  Mosses  of  North  America  (1884),  p. 
285,  we  find  not  a  single  positive  or  new  character  as- 


Holzinger:    MUSCI  OF  THE  INTERNATIONAL  BOUNDARY.       47 

signed  to  Homalia  jainesii,  except  leaves  "  striolate 
lengthwise  when  dry."  And  this  point  is  not  borne  out 
by  the  actual  examination  of  James'  own  material. 

It  appears,  therefore,  that  Homalia  jamesii  is  too  close 
tO-ff.  trichomanoidcs ;  that  Homalia  macounii  is  identical 
with  Homalia  jamesii;  that  the  only  difference  is  found 
in  the  more  obtuse  leaves  of  our  species,  which  proves  to 
be  a  variable  character,  and  therefore  that  it  should  not 
stand  as  a  distinct  species,  hardly  deserving  the  name  of 
a  variety.  As  a  variety  it  must  be  called : 

Homalia  trichomanoides  jamesii  (SCHIMP.). 

H.  jamesii  Schimp.  in  Syn.  (  ),  p.  473. 

H.  macounii  in  Mac.  Cat.  (1892),  p.  163. 
The  geographical  distribution  of  this  variety  of  Homalia 
trichomanoidcs,  includes  necessarily  all  the  stations 
cited  in  Macoun's  Catalogue  for  H.  macounii  with 
those  given  for  H.  jamesii  in  Lesquereux  and  James' 
Manual. 

60.  Myurella  careyana  SULL. 

On  Grand  Portage  island,  north  shore  of  Lake  Superior 
(June  23,  1897). 

61.  Myurella  julacea  SCH. 

At  the  base  of  Kawasatchong  falls,  shore  of  Fall  lake, 

near  Camp  i  (June  8-10,  1897). 

On  Grand  Portage  island,  north  shore  of  Lake  Superior 
(June  23,  1897). 

62.  Leskea  nervosa  MYR. 

On  the  Prairie  portage,  shore  of  Basswood  lake,  near  the 
rapids  from  Sucker  lake  (June  12,  13,  1897). 

63.  Leskea  polycarpa  EHRH. 

At  the  farther  end  of  the  portage  across  the  United 
States  peninsula,  shore  of  Basswood  lake  (June  12, 
1897). 

On  the  Prairie  portage,  shore  of  Basswood  lake,  near  the 
rapids  from  Sucker  lake  (June  12,  13,  1897). 

64.  Leskea  polycarpa  paludosa  SCH. 

On  the  way  from  Ely  to  Winton,  shore  of  Fall  lake 
(JuneS,  1897). 


48  MINNESOTA    BOTANICAL    STUDIES. 

Along  the  shore  of  Fall  lake,  near  Camp  i  (June  8-10, 
1897),  abundant. 

65.  Anomodon  attenuatus  HARTM. 

Shore  of  Fall  lake,  near  Camp  i  (June  8-10,  1897). 
On  a  small  point  of  land,  'at  the  base  of  the  United  States 

peninsula,  shore  of  Basswood  lake  (June  n,  1897). 
On  the  Prairie  portage,  shore  of  Basswood  lake,  near  the 

rapids  from  Sucker  lake  (June  12,  13,  1897). 

66.  Anomodon  minor  (P.  BEAUV.)  FURN. 

On  the  shore  of  Fall  lake,  base  of  Kawasatchong  falls, 
near  Camp  i  (June  8-10,  1897). 

67.  Anomodon  rostratus  SCH. 

On  the  shore  of  Fall  lake,  base  of  Kawasatchong  falls, 

near  Camp  i  (June8-io,  1897). 
On  a  small  point  of  land,  at  the  base  of  the  United  States 

peninsula,  shore  of  Basswood  lake  (June  n,  1897). 
On  the  Prairie  portage,  shore  of  Basswood  lake,  near  the 

rapids  from  Sucker  lake  (June  12,  13,  1897). 

68.  Pylaisia  heteromalla  SCH. 

On  trees  along  the  shore  of  Fall  lake,  near  Camp  i  (June 
8-10,  1897). 

On  a  small  point,  at  the  base  of  the  United  States  penin- 
sula, Basswood  lake  (June  u,  1897). 

69.  Pylaisia  polyantha  SCH. 

On  trees  along  the  shore  of  Fall  lake,  near  Camp  i  (June 

8-10,   1897). 

On  the  Prairie  portage,  shore  of  Basswood  lake,  near 
the  rapids  from  Sucker  lake  (June  12,  13.  1897). 

70.  Platygyrium  repens  SCH. 

Shore  of  Fall  lake,  near  Camp  i.  On  dead  logs  (June 
8-10,  1897). 

71.  Cylindrotheciurn  seductrix  SULLIV. 

Same  locality  as  above. 

72.  Climacium  americanum  BRID. 

Same  locality  as  above. 

Portage  from  Mountain  lake  to  Moose  lake  (June  20, 
1897). 


Holzinger :    MUSCI  OF  THE  INTERNATIONAL  BOUNDARY.       49 

73.  Thuidium  abietinum  Sen. 

Shore  of  Fall  lake,  near  Camp   i   (June  8-10,  1897). 
On  the  Prairie  portage,  shore  of  Basswood  lake   (June 

12,  13,  1897). 
On  the  portage  from   South-Fowl  lake  to  Pigeon  river 

(June  21,  1897). 
On  Grand  Portage  island,  north  shore  of    Lake  Superior 

(June  23,  1894). 

74.  Thuidium  recognitum  LINDB. 

Shore  of  Fall  lake,  near  Camp  i  (June  8-10,  1897). 

75.  Thuidium  philiberti  LIMPR. 

At  camp,  shore  of  Fall  lake  (June  8-10,  1897). 

76.  Brachythecium  campestre  SCH. 

Shore    of  Fall  lake  near  Camp  i   (June  8-10,  1897). 

77.  Brachythecium  flexicaule  REN.  and  CARD. 

On  the  Prairie  portage,  shore  of  Basswood  lake  (June 

12,   13,   1897). 

78.  Brachythecium  oxycladon  (BRID.).     Grout. 

At   the   base  of   the  United  States  peninsula,  shore  of 

Basswood  lake  (June  n,  1897). 
On  the  Prairie  portage,  shore  of   Basswood  lake  (June 

12,   13,   1897). 

On  Grand  Portage  island,  north  shore  of  Lake  Superior 

(June  23,  1897). 
This  last  is  pronounced  a  "  slender  form  "  by  Mr.  Grout. 

Brachythecium  plumosum  SCH.? 

On  the  Prairie  portage,  shore  of   Basswood  lake  (June 

12,    13,    1897). 

80.  Brachythecium  salebrosum  SCH. 

Shore  of  Fall  lake,  near  Camp  i   (June  8-10,  1897). 

81.  Brachythecium  starkei  SCH. 

At  the  base  of    the   United   States   peninsula,  shore  of 
Basswood  lake  (June  n,  1897). 

82.  Eurynchium  robustum  (ROELL.). 

At  the  base  of   the  United  States    peninsula,  shore  of 

Basswood  lake  (June  n,  1897). 
On   Basswood  lake,   at  the  farther  end   of  the  portage 

across  the  United  States  peninsula  (June  12,  1897). 


50  MINNESOTA    BOTANICAL    STUDIES. 

Mr.  Grout  remarks  that  these  plants  vary  toward  E. 
strigosum ;  they  are  probably  only  large  forms  of  this 
species. 

83.  Eurynchium  strigosum  SCH. 

Shore  of  Fall  lake,  near  Camp  i  (June  8-10,  1897). 
On  the  portage  from  South  lake  to  Rat  lake  ( June  20, 

1897). 
On  the  portage  from  South-Fowl  lake  to  Pigeon  river 

(June  21,  1897). 
On  Grand  Portage  island,  north  shore  of  Lake  Superior 

June  23,  1897). 

84.  Raphidostegium  recurvans  L.  and  J. 

Shore  of  Fall  lake,  near  Camp  i  (June  8-10,  1897). 
On  the  point  of  land  at  the  base  of  the  United  States 

peninsula,  Basswood  lake  (June  u,  1897). 
On  Safety  island,  Lake  Saganaga  (June  16,  1897). 

85.  Plagiothecium  denticulatum  SCH. 

On  the    road  from  Ely  to  Winton,  shore  of    Fall  lake 

(JuneS,  1897). 
At  Camp    i,  shore  of   Fall   lake,    near    Kawasatchong 

falls  (June  8-10,  1897). 

86.  Plagiothecium  muhlenbeckii  SCH. 

At  Camp  i,  shore  of  Fall  lake  (June  8-10,  1897). 

87.  Plagiothecium  sylvaticum  SCH. 

At  Camp  i,  shore  of  Fall  lake  (June  8-10,  1897). 
On  Grand  Portage  island,  north  shore  of  Lake  Superior 
(June  23,  1897). 

88.  Amblystegium  adnatum  L.  and  J. 

At   Camp    i,   shore    of    Fall    lake  (June   8-10,    1897). 
Det.  by  G.  N.  Best. 

89.  Amblystegium  serpens  SCH. 

On   the    portage    from    Mountain    lake    to    Moose    lake 
(June  20,  1897). 

90.  Hypnum  chrysophyllum  BRID. 

At  Camp  i,  shore  of  Fall  lake  (June  8-10,  1897). 

91.  Hypnum  cupressiforme  ericetorum  B.  S. 

At  Camp  i,  shore  of  Fall  lake  (June  8-10,  1897). 
At   the  lower   end    of    Pipestone    rapids,  on    Basswood 
lake  (June  10,  n,  1897). 


Holzinger :   MUSCI  OF  THE  INTERNATIONAL  BOUNDARY.       51 

92.  Hypnum  crista-castrensis  L. 

On  the  road  from  Ely  to  Winton,  on  Fall  lake  (June  8, 

1897). 

At  Camp  i,  on  Fall  lake  (June  8-10,  1897). 
At   the    lower  end   of  Pipestone    rapids,   on    Basswood 

lake  (June  10,  u,  1897). 
At  the  base  of  the  United  States  peninsula,  Basswood 

lake  (June  n,  1897). 
Near  Camp  3,  at  the  farther  end  of  the  portage  across 

the  United  States  peninsula,  on  Basswood  lake  (June 

12,  1897). 
On  Prairie  portage,  shore  of  Basswood  lake  (June  12,  13, 

1897). 
On  Safety  island,  Lake  Saganaga  (June  16,  1897). 

93.  Hypnum  filicinum  trichodes  BRID. 

On  Grand  Portage  island,  north  shore  of  Lake  Superior 

(June  23,  1897). 
Dr.  Best  remarks  that  this  approaches  the  variety  acicu- 

linum  C.  M.  and  K. 

94.  Hypnum  haldanianum  GREV. 

On  the  road  from  Ely  to  Fall  lake  (June  8,  1897). 

At    Camp   i,   on   Fall    lake,    near   Kawasatchong  falls 

(June  8-10,  1897). 
At  the  lower  end  of  the  Pipestone   rapids,  on  Basswood 

lake  (June  10,  u,  1897). 
At  the  base  of  the  United  States  peninsula,  on  Basswood 

lake  (June  u,  1897). 
On  the  portage  from  South  lake  to  Rat  lake  (June  20, 

1897). 

95.  Hypnum  hispidulum  BRID. 

On  the  road  from  Ely  to  Fall  lake  (June  8,  1897). 
At  Camp  i,  on  Fall  lake  (June  8-10,  1897). 

96.  Hypnum  reptile  RICH. 

At  Camp  i,  on  Fall  lake  (June  8-10,  1897.) 
At  a  small  point  of  land  at  the  base  of  the  United  States 
peninsula,  Basswood  lake  (June  n,  1897). 

97.  Hypnum  schreberi  WILLD. 

At  Camp  i,  on  Fall  lake  (June  8-10,  1897). 
At  the  lower  end  of  the  Pipestone  rapids,  on  Basswood 
lake,  near  Camp  2  (June  10,  n,  1897). 


52  MINNESOTA    BOTANICAL    STUDIES. 

At  the  base  of  the  United  States  peninsula,  Basswood 

lake  (June  u,  1897). 
At  the  farther  end  of  the  portage  across  the  United  States 

peninsula,  Basswood  lake  (June  12,  1897). 
On  Safety  island,  Lake  Saganaga  (June  16,  1897). 
At  the  east  end  of  Gunflint  lake  (June  20,  1897). 
On  Grand  Portage  island,  north  shore  of  Lake  Superior 

(June  23,  1897). 

98.  Hypnum  uncinatum  HEDW. 

On  the  road  from  Ely  to  Fall  lake  (June  8,  1897). 
On  the  portage  across  the  divide  (June  20,  1897). 

99.  Holocomium  splendens  SCH. 

At  Camp  i,  shore  of  Fall  lake  (June  8-10,  1897). 

At  the  base  of  the  United  States  peninsula,  Basswood 

lake  (June  n,  1897). 
On  the  Prairie  portage,  shore  of  Basswood  lake  (June  12, 

i3.  l897)- 
On  Safety  island,  Lake  Saganaga  (June  16,  1897). 

At  the  east  end  of  Gunflint  lake  (June  10,  1897). 
On  Grand  Portage  island,  north  shore  of  Lake  Superior 
(June  23,  1897). 

100.  Hylocomium  triquetrum  SCH. 

On  the  road  from  Ely  to  Fall  lake  (June  8,  1897). 
At  Camp  i,  shore  of  Fall  lake  (June  8-10,  1897). 
At  the  farther  end  of  the  portage  across  the  United 

States  peninsula,  shore  of  Basswood  lake   (June  12, 

1897). 

On  Safety  island,  Lake  Saganaga  (June  16,  1897). 
At  the  east  end  of  Gunflint  lake  (June  20,  1897). 
On  Grand  Portage  island,  north  shore  of  Lake  Superior 

(June  23,  1897). 


VII.     THE  INFLUENCE  OF  GASES  AND   VAPORS 
UPON  THE  GROWTH  OF  PLANTS. 


EMIL  P.  SANDSTEN. 


INTRODUCTION. 

In  recent  years  considerable  attention  has  been  paid  by  phys- 
iologists to  the  influence  of  various  chemical  agents  upon  the 
growth  of  plants,  and  the  results  thus  far  obtained  seemed  to 
warrant  further  investigation  along  this  line.  The  work  here- 
tofore has  been  confined  almost  exclusively  to  the  lower  plant 
forms,  which  are  more  easily  attacked  by  the  difficult  technique 
which  is  bound  up  in  this  kind  of  inquiries.  The  recent  prelim- 
inary results  of  Johanssen  (i)  were  announced  shortly  after  this 
work  was  begun,  and  it  was  thought  advisable  to  extend  the 
work  to  cover  the  phases  of  the  vegetative  period  as  well  as  rest- 
ing seeds,  etc.  To  some  extent  the  writer  has  had  in  mind  the 
ultimate  application  of  the  reactions  obtained  in  practical  garden- 
ing though  such  results  are  reserved  for  verification  and  further 
trial.  The  work  was  done  during  the  fall  and  winter  of  1897 
and  1898  in  the  laboratories  of  plant  physiology  in  the  Univer- 
sity of  Minnesota  under  the  direction  of  Dr.  D.  T.  Mac  Dougal, 
to  whom  the  writer  is  greatly  indebted  for  his  valuable  advice 
and  kind  criticism. 

MATERIAL  AND  METHODS. 
The  experiments  may  be  conveniently  classified  as  follows : 

1.  The  influence  of  gases  and  vapors  upon  seeds. 

2.  The  influence  of  gases  and  vapors  upon  seedlings. 

3.  The  influence   of  gases  and  vapors  upon  growing  shoots. 

4.  The  influence  of  gases  and  vapors  upon    resting   bulbs, 
corms,  etc. 

5.  The  influence  of  gases  and  vapors  upon  plants  growing  in 
water  cultures. 

The  reagents  used  were  alcohol,  ammonia,  carbon  bisulphide, 


54  MINNESOTA    BOTANICAL    STUDIES. 

chloroform,  ether,  nitrous  oxide  and  oxygen.  Small  quantities 
of  alcohol  (methyl),  ammonia  (hydrate),  carbon  bisulphide,  ether 
and  chloroform  were  placed  in  tubes  inside  of  closed  receivers 
and  allowed  to  vaporize  into  the  air  enclosed.  The  nitrous  ox- 
ide was  the  commercial  mixture,  N2O  90  parts,  N  8.86,  O  1.13. 
In  certain  experiments  the  pure  gas  which  had  been  obtained 
from  ammonium  nitrate  was  used.  Commercial  oxygen  from 
tanks  was  used. 

As  a  means  of  control  and  test  of  the  actual  efficiency  of  the 
reagents,  leaves  of  Philotria  (Elodea)  and  hairs  of  Tradescantia, 
Tomato,  Begonia,  Pelargonium  and  Geranium  were  mounted 
in  an  Engelmann  gas  chamber  and  subjected  to  their  action. 
These  tests  were  carried  on  at  a  room  temperature  of  16  to  23°  C., 
and  the  results  noted  below  are  quite  in  harmony  with  those 
given  by  previous  writers. 

Oxygen.  The  movements  of  protoplasm  are  greatly  acceler- 
ated in  an  atmosphere  of  free  oxygen  for  five  to  seven  minutes, 
after  which  the  movements  gradually  diminish  until  they  cease 
entirely.  If  the  living  cell  is  kept  under  the  influence  of  free  oxy- 
gen for  considerable  length  of  time  it  dies,  but  no  apparent  change 
in  the  structure  or  behavior  of  the  protoplasm  could  be  noticed. 

Nitrous  oxide  (N2O).  This  gas  has  the  same  general  effect 
on  living  protoplasm  as  oxygen  with  the  exception  that  it  is  less 
and  does  not  kill  the  cell  even  when  exposed  for  several  days 
in  an  atmosphere  of  90  per  cent,  nitrous  oxide.  The  duration  of 
active  movements  varies  from  three  to  five  minutes  (Moeller  V.). 

Chloroform  and  Ether.  The  action  of  these  two  reagents 
are  about  the  same.  Aqueous  solutions  containing  1/20000  part 
of  reagents  at  first  slightly  increase  the  movements  of  the  proto- 
plasm. By  increasing  the  strength  of  the  solution  the  rapidity 
of  movement  was  also  increased,  but  the  reaction  time  was  very 
much  shortened.  Strong  solution  causes  vacuolization  and  par- 
alyzes the  protoplasm. 

Ammonia.  Weak  aqueous  solution  containing  from  1/30000 
to  1/20000  parts  of  ammonia  does  not  seem  to  modify  the  activity 
of  the  protoplasm  when  subjected  to  its  action  for  a  short  time 
only.  Stronger  solution  produces  vacuolization  and  slightly  ac- 
celerates the  movements  of  the  protoplasm  for  a  minute  or  two. 

Carbon  bisulphide. — The  smallest  possible  quantity  that  could 
be  introduced  arrested  all  movements. 

Alcohol. — Aqueous  solutions  containing  1/20000  to  i/ioooo 
parts  of  alcohol  had  no  visible  effect  upon  the  protoplasm.  A 


Sandstcn :  INFLUENCE  OF  GASES  AND  VAPORS  UPON  GROWTH.    55 

2  per  cent,  solution  excited  rapid  irregular  movements  which 
stopped  inside  of  two  minutes.  Vacuolization  followed  rapidly 
and  the  cell  was  killed  inside  of  ten  minutes. 

In  the  experiments  where  seeds  and  seedlings  were  used,  Zca 
mat's,  Vicia  and  Phaseolus  were  employed  exclusively.  Straw- 
berry plants  of  the  common  cultivated  kind  were  used  in  the  ex- 
periments with  growing  shoot  and  proved  well  adapted  to  the 
work.  The  strawberry  plants  were  taken  from  the  bed  on  No- 
vember 6,  1897,  and  carefully  selected  with  reference  to  vigor 
and  equality.  Two  lots  of  plants  were  selected,  one  lot  com- 
posed of  plants  one  season  old,  the  other  composed  of  plants  two 
seasons  old.  The  plants  were  placed  in  three-  and  four-inch 
pots  respectively  November  9th,  and  set  in  a  cold  frame  where 
they  remained  until  December  9th,  when  they  were  taken  to  the 
green  house  and  put  under  the  experimental  conditions  described 
below. 

Dormant  bulbs  and  corms  of  Ariscema,  Narcissus,  Hyacinth, 
Tulip,  Frcesia  and  Crocus  were  used  for  material  in  the  resting 
stage. 

In  the  experiments  with  gases  in  nutrient  solution  in  water 
culture  seedlings  of  Zea  mats  were  used.  A  large  number  of 
seeds  were  germinated  in  clean  saw-dust  and  when  the  seedlings 
had  attained  the  desired  growth  the  specimens  which  were  to 
be  used  in  the  experiment  were  carefully  selected  for  vigor  and 
equality.  The  vessels  holding  the  nutrient  solution  were  glazed 
earthen  jars  of  two  litres  capacity.  The  tops  of  the  earthen  jars 
were  fitted  with  covers  made  of  plaster  of  paris.  Through  each 
cover  two  holes  were  grilled,  one  for  the  seedlings  and  a  second 
to  admit  the  necks  of  inverted  flasks  of  gas.  The  seedlings 
were  fastened  in  the  openings  in  the  covers  by  means  of  a  per- 
forated cork  after  the  usual  manner  in  water  cultures.  The 
flasks  were  filled  with  water,  inverted  with  the  necks  immersed 
in  the  culture  fluid  and  filled  with  gas  by  displacement  through 
a  bent  glass  delivery  tube. 

The  following  formula  was  used  in  making  up  the  nutrient 
solutions  : 

Potassium   nitrate 25-    Sm- 

Sodium  chloride 12.5    " 

Calcium   sulphate 12-5    " 

Magnesium    sulphate I2-5 

Calcium  phosphate I2-5 

Water...  250.    cc. 


56  MINNESOTA    BOTANICAL    STUDIES. 

The  solution  was  diluted  to  12500  cc.  and  from  five  to  eight 
drops  of  ferric  chloride  were  introduced  in  each  jar  before 
using. 

The  bell  jars  used  in  enclosing  the  bulbs  and  shoots  had 
ground  edges  and  were  set  upon  ground  glass  plates  which 
had  been  anointed  with  a  preparation  composed  of  vaseline, 
tallow  and  resin,  to  make  the  connection  absolutely  air  tight. 
The  temperature  was  kept  as  constant  as  possible. 

The  pressure  of  one  atmosphere  is  to  be  understood  where 
not  otherwise  stated. 

i.    THE  INFLUENCE  OF  GASES  AND  VAPORS  UPON  THE  GER- 
MINATION OF  SEEDS. 

A.  Gases.    The  gases  used  were  nitrous  oxide  and  oxygen. 
Seeds  of  Phaseolus  multiflorus  and   Vicia  faba  were  soaked  in 
water  for  twenty-four  hours  and  from  these  were  selected  ten 
normal    specimens    for    each    experiment.     They   were    then 
placed  on  sections  of  cork,  which  had  previously  been  soaked 
in  water  and  introduced  into  the  bell  jar  under  water  so  as  to 
prevent  any  air  from  gaining  admittance.     Duplicate    exper- 
iments and   duplicate   controls  were  set  up.     The  capacity  of 
bell  jars  was  two  litres. 

The  results  obtained  with  nitrous  oxide  and  oxygen  agree 
with  previous  experiments  in  the  same  line.  (Detmer  II.)  The 
seed  germinated  readily  in  an  atmosphere  of  free  oxygen,  but 
failed  to  do  so  in  an  atmosphere  of  nitrous  oxide.  The  nitrous 
oxide  gas  did  not  kill  the  seeds,  as  they  afterwards  germinated 
under  a  bell  jar  in  ordinary  air.  The  N2O  used  here  was  ob- 
tained from  ammonium  nitrate. 

B.  Valors.     Seeds  of  Phaseolus  mufajftorusiand   Vicia  faba 
were  placed  under  bell  jars,  4000  cc.  capacity,  tightly  secured 
to  glass  plates.     Twelve  dry  seeds  of  each  kind  were  placed 
under  each  bell  jar,  together  with  a  small  glass  vial  containing 
accurately    measured    quantities    of   the    reagent.     The    seeds 
were  kept  under  the  bell  jars  for  nine  days,  when  they  were 
taken  out  and  each  lot  planted   separately  in  four-inch  pots. 
The  control  experiment  was  treated  exactly  in  the  same  way  as 
the  others  with  the  exception  of  the  omission  of  the  chemicals. 
The  plants  were  growing  side  by  side  and  received  the   same 
treatment. 


Sandsten:  INFLUENCE  OF  GASES  AND  VAPORS  UPON  GROWTH.   57 

TABLE  I. 
(KEY.) 

I.    1/32000  parts  of  NOH4 

II.    1/28000     "     "        " 

III.  1/24000     "     "        " 

IV.  1/20000       "       "  " 

(a)  Phaseolus  multiflorus. 

(b)  Viciafaba. 

(c)  Control  of  Phaseolus. 

(d)  Control  of    Vicia. 


Per  cent,  of 
germination. 

Average  height 
10  days 
after  planting. 

Average  height 
28  days 
after  planting. 

Time  of 
blooming. 

(a) 
j    (b) 
(c) 
(d) 

IOG 

90 

IOO 
IOO 

19  mm. 
33  mm. 
-(-(a)  51  mm. 
•f  (b)  59  mm. 

28      cm. 
12.5  cm. 
30.5  cm. 
15      cm. 

30  days. 

33  days. 

(a) 

II    (b) 
IL    (c) 

(d) 

IOO 
2 
IOO 
IOO 

18.5  mm. 
25      mm. 
+  (a)  51      mm. 
+  (b)  59      mm. 

22.5  cm. 
*n.5  cm. 
30.5  cm. 
15      cm. 

29  days. 

33  days. 

(a) 
HI.   $ 
(d) 

So 
20 

IOO 
IOO 

30  mm. 
51  mm. 
4-(a)  51  mm. 
+  (b)  59  mm. 

27      cm. 
11.5  cm. 
30.5  cm. 
15      cm. 

31  days. 

33  days. 

(a) 

iv   (b> 

(c) 
(d) 

0 

o 

IOO 
IOO 

0 

o 
+  (a)  51  m. 
+  (b)  59  m. 

o 
o 
30.5  cm. 
15      cm. 

33  days. 

It  will  be  seen  that  very  small  quantities  of  ammonia  vapors  are 
not  fatal  to  the  germination.  In  none  of  the  experiments  had 
any  of  the  seeds  germinated  during  the  nine  days  they  were 
under  the  bell  jars,  nor  had  any  of  the  seeds  in  the  control  ger- 
minated. The  odor  of  ammonia  from  the  seeds  treated  could 
readily  be  detected.  It  was  noticed  throughout  the  experiments 
that  the  plants  from  the  treated  seeds  had  a  deeper  green  color 
than  the  control.  This  was  especially  noticeable  in  the  case  of 
a  and  b  in  series  I  and  II.  Nor  did  it  appear  that  the  ammonia 
vapor  had  any  subsequent  bad  effect  on  the  plants ;  on  the  con- 


One  plant  only. 


58  MINNESOTA    BOTANICAL    STUDIES. 

trary  in  series  I  and  II  it  seems  to  have  hastened  the  time  of  the 
flowering  by  three  to  four  days.  Vicia  faba  is  more  suscep- 
tible to  ammonia  vapors  than  Phaseolus  multiflorus.  As  in  the 
case  of  Phaseolus  multiflorus  the  leaves  were  darker  than  in  the 
control.  The  measurements  given  in  the  table  above  represent 
the  average  growth  of  shoot  of  twelve  plants. 

2.  THE  INFLUENCE  OF  GASES  AND  VAPORS  UPON  SEEDLINGS. 

A.  Gases. — Nitrous  oxide  of  oxygen   and  seedling    of  Zca 
mais  and  Phaseolus  multiflorus  were  employed  in  these  experi- 
ments.    The  seedlings  were    carefully    measured  and  placed 
under  bell  jars  while  full  of  water  which  was  displaced  by  the 
gases. 

The  following  quantities  of  gases  were  used  :  400  cc.  of  ni- 
trous oxide  in  2000  cc.  of  air  and  an  atmosphere  of  free  nitrous 
oxide ;  400  cc.  of  oxygen  in  2000  cc.  of  air  and  an  atmosphere 
of  free  oxygen.  These  were  set  up  in  duplicates  and  the  con- 
trol was  also  in  duplicate.  The  results  of  the  experiments 
showed  an  increase  in  growth  for  seedlings  in  the  two  oxygen 
experiments  and  also  for  the  nitrous  oxide  experiment  in  which 
400  cc.  in  2000  cc.  of  air  was  used.  The  average  increase  in 
the  two  oxygen  experiments  and  the  control  for  24  hours 
was  little  less  than  8  mm.  The  seedlings  in  the  atmosphere  of 
free  oxygen  did  not  average  as  much  as  those  in  the  partial  at- 
mosphere of  oxygen.  The  average  was  5  mm. 

The  seedlings  in  the  experiments  in  which  400  cc.  in  2000 
cc.  of  air  was  used  showed  a  slight  increase  in  growth  over 
the  control,  amounting  on  an  average  to  3  mm.  The  seedlings  in 
an  atmosphere  of  free  nitrous  oxide  did  not  make  any  growth, 
but  were  alive  when  taken  from  the  bell  jar.  The  temperature 
during  the  time  the  experiments  were  running  varied  from  21- 
23°  C. 

B.  Vapors. — The   following    chemicals  were  used :    Ether, 
chloroform,  carbon  bisulphide,  alcohol  and  ammonia.      Seeds 
of  Zea  mais  were  germinated  in  clean  saw-dust  and  when  the 
roots  had  attained  a  length  of  15  to  20  mm.   and  the  plumule 
from  10  to   15  mm.  a  uniform  lot  was  selected  for  the  experi- 
ments.   The    roots    and    shoots  were    carefully  measured    and 
marked  with  India  ink.     The  seedlings  were  next  placed  under 
the  bell  jars  of  2000  cc.  capacity  upon  moist  saw-dust.     The 
chemicals  were  accurately  measured  out  and  put  into  small  glass 


Sandsten :  INFLUENCE  OF  GASES  AND  VAPORS  UPON  GROWTH.   59 

bottles  containing  100  cc.  of  water  and  then  placed  under  the 
bell  jars  with  the  seedlings.  The  temperature  during  the  ex- 
periments varied  from  21-23°  C.  A  new  quantity  of  chemical 
was  introduced  each  time  after  the  seedlings  were  measured, 
thus  keeping  the  amount  of  vapors  constant  throughout  the  time 
the  experiments  were  running.  Commercial  mixtures  of  nitrous 
oxide  were  used  in  the  above  series. 

The  results  are  given  below  in  Tables  II  and  III. 

TABLE  II. 

I.  .2  cc.  of  ether  in  2000  cc.  of  air. 

II.  .2  cc.  of  chloroform  in  2000  cc.  of  air. 

III.  .2  cc.  of  carbon  bisulphide  in  2000  cc.  of  air. 

IV.  .5  cc.  alcohol  in  2000  cc.  of  air. 
V.  Control. 

a.  Plumule. 

b.  Root. 


Series. 

I/> 

c 
Jt 

HH 

Average 
growth  of 
plumule  and 
root  of  10 
seedlings  in 
3  hours. 

Average 
growth  of 
plumule  and 
root  of  10 
seedlings  in 
5  hours. 

Average 
growth  of 
plumule  and 
root  of  10 
seedlings  in 
24  hours. 

Average 
growth  of 
plumule  and 
root  of  10 
seedlings  in 
48  hours. 

Total  gain 
or  loss  over 
the  control. 

I. 

a. 
b. 

1.4    mm. 
1.75  mm. 

2.15  mm. 
3-5    mm- 

9.75  mm. 
26       mm. 

28.25  mm. 
39.5    mm. 

+  2.45  mm. 
+  7.5    mm. 

II. 

a. 
b. 

1.25  mm. 
1.4    mm. 

2.25  mm. 
2.5    mm. 

20.3    mm. 
28.66  mm. 

32.66  mm. 
37.66  mm. 

4-  6.66  mm. 
-(-  5-66  mm. 

III. 

a. 

b. 

1.4  mm. 
I.     mm. 

2.4    mm. 
2.88  mm. 

7.25  mm. 
6.5    mm. 

14.25  mm. 
9.      mm. 

—  10.55  mm. 
—  28.      mm. 

IV. 

a. 

b. 

1.5  mm. 
2.    mm. 

2.75  mm. 
2.5    mm. 

14.25  mm. 
8.5    mm. 

28.5    mm. 
18.25  mm- 

—  13.75  mm. 

V. 

a. 

b. 

1.2    mm. 
1.45  mm. 

2.1  mm. 
2.5  mm. 

ii.  8  mm. 
16.5  mm. 

25.8  mm. 
32.     mm. 

60 


MINNESOTA    BOTANICAL    STUDIES. 


TABLE  III. 

I.     .4  cc.  of  ether  in  2000  cc.  of  air. 
II.     .4  cc.  of  chloroform  in  2000  cc.  of  air 

III.  .4  cc.  of  carbon  bisulphide  in  2000  cc.  ot  air. 

IV.  I.     cc.  of  alcohol  in  2000  cc.  of  air. 
V.      .2  cc.  of  ammonia  in  2000  cc.  of  air. 

VI.  Control. 


Series. 

Roots. 

Average  growth 
of  roots  of  10 
seedlings  for 
6  hours. 

Average  growth 
of  roots  of  10 
seedlings  for 
24  hours. 

Total  loss  or 
gain  over  the 
control. 

I. 

1.3  mm. 

10.     mm. 

-   3.1  mm. 

II. 

I.     mm. 

7.5  mm. 

-    5.6  mm. 

III. 

.6  mm. 

.6  mm. 

—  12.5  mm. 

IV. 

dead. 

dea^l. 

V. 

1.3  mm. 

13.2  mm. 

-+-      .1  mm. 

VI. 

1.17  mm. 

13.1  mm. 

From  the  above  tables  it  will  be  seen  that  a  very  small  amount 
of  carbon  bisulphide  or  ammonia  vapors  is  very  injurious  to 
young  seedlings,  while  ether,  chloroform  and  alcohol  vapors 
in  minute  quantities  are  not  injurious  when  the  plant  is  not 
subjected  to  their  prolonged  action.  On  the  contrary,  small 
amounts  of  ether  and  chloroform  vapors  seem  to  accelerate 
growth. 

3.  THE  INFLUENCE  OF   GASES  AND  VAPORS  UPON    GROWING 

SHOOTS. 

A.  Gases. — Nitrous  oxide  and  oxygen  were  used  in  these 
experiments  in  the  following  quantities:  25  per  cent.,  50  per 
cent,  and  100  per  cent.  The  plants  were  kept  under  the  bell 
jars  for  twenty  days. 


Sandsten  :  INFLUENCE  OF  GASES  AND  VAPORS  UPON  GROWTH.    61 


TAI5LE    IV. 

I.   25  per  cent,  of  gas  in  4000  cc.  of  air. 
II.   50  per  cent,  of  gas  in  4000  cc.  of  air. 

III.  One  atmosphere  of  free  gas. 

IV.  Control. 

a.  Nitrous  oxide. 

b.  Oxygen. 


Series. 

Chem- 
icals. 

Time  of  first 
flowers. 

Number  of 
flowers. 

Number  of 
leaves. 

Scale  of  vigor. 
Control  taken 
as  standard. 

T 

a. 

43  days. 

3 

8 

112 

b. 

64  days. 

3 

7 

103 

II. 

a. 
b. 

28  days, 
no  flowers. 

17 
no  flowers. 

H 
3 

125 
45 

a. 

42  days. 

3 

7 

IO2 

b. 

dead. 

IV. 

a. 

b. 

44  days. 
46  days. 

i 
3 

6 

7 

IOO 

too 

In  the  above  table  the  plants  treated  with  nitrous  oxide  show 
a  marked  increase  in  vigor  and  flowering  capacity.  The  leaves 
were  of  a  dark  green  color  and  very  large.  The  leaf  petioles 
were  somewhat  shortened,  giving  the  plants  a  stocky  appear- 
ance. The  root  systems  of  the  plants  treated  with  nitrous  ox- 
ide were  very  strong.  All  evidence  seems  to  point  to  the  con- 
clusion that  the  treatment  was  beneficial  to  the  plants.  The 
oxygen  also  appeared  to  be  beneficial  to  the  plants  when  used 
in  quantities  not  exceeding  50  per  cent.  In  an  atmosphere  of 
free  oxygen  the  plants  showed  no  deviation  from  the  normal 
while  in  the  gas,  but  upon  the  removal  of  the  bell  jar  the  plants 
soon  began  to  show  signs  of  decay.  The  plants  treated  with 
oxygen  exhibited  a  marked  elongation  of  the  petioles. 

B.  Vapors. — Ammonia  and  chloroform  were  used  in  these 
experiments  in  the  following  quantities  :  i/ioooo,  1/15000  and 
i  40000  parts.  The  capacity  of  the  bell  jars  was  7500  cc.  The 
reagents  were  introduced  in  an  aqueous  solution  of  100  cc.  The 
plants  were  kept  under  the  bell  jars  for  26  hours.  Upon  exam- 
ination it  was  found  that  the  plants  which  had  been  subjected 
to  the  influence  of  i/ioooo  part  of  ammonia  or  chloroform  vapors 


62  MINNESOTA    BOTANICAL    STUDIES. 

were  dead.  The  leaves  had  assumed  a  dirty  brown  color.  The 
center  of  the  shoot  was  badly  discolored.  The  appearance  of 
the  plants  was  very  similar  to  that  of  a  frozen  plant. 
The  plants  which  were  subjected  to  1/15000  part  of  the  reagents 
were  badly  effected,  the  outer  leaves  were  dark  brown  but  the 
center  was  not  affected.  The  plants  grew  but  remained  weak 
and  straggling  throughout  the  time  the  experiments  were  run- 
ning. The  action  of  the  two  reagents  seemed  to  be  the  same, 
little  or  no  difference  could  be  detected. 

The  plants  which  were  kept  in  an  atmosphere  containing 
1/40000  part  of  the  reagents  did  not  appear  to  be  visibly  affected 
when  taken  from  the  bell  jars.  The  subsequent  influence  of  the 
reagents  was,  however,  very  marked,  especially  on  the  plants 
in  the  chloroform  experiment.  Compared  with  the  control 
plants  at  the  end  of  the  experiment,  February  10,  with  which 
they  were  equal  at  the  start,  they  showed  a  great  advance. 

4.  THE  INFLUENCES  OF  GASES  AND  VAPORS  UPON  RESTING 

BULBS. 

The  bulbs  used  in  these  experiments  were  Ariscema  triphyl- 
lum,  Narcissus,  Hyacinth,  Crocus,  and  Freesia.  The  reagents 
used  were  oxygen,  nitrous  oxide,  ether,  chloroform,  carbon  bi- 
sulphide, ammonia  and  alcohol.  These  experiments  were  started 
in  November  and  December,  months  in  which  bulbs  of  this  kind 
are  very  hard  to  start,  since  they  require  a  certain  period  of  rest 
before  beginning  growth  and  this  period  generally  extends 
through  the  months  of  October,  November  and  December. 

The  bulbs  were  kept  under  bell  jars  and  the  reagents  which 
were  in  a  liquid  form  were  introduced  in  aqueous  solution. 
Where  gases  were  used  in  the  experiments  they  were  intro- 
duced by  displacement. 

A.  Gases. — Narcissus  bulbs  were  placed  under  bell  jars  con- 
taining oxygen  in  the  following  proportion:  20  per  cent.,  50 
per  cent,  and  100  per  cent.  The  capacity  of  the  bell  jars  was 
4000  cc. 


Sandsten  :  INFLUENCE  OF  GASES  AXD  VAPORS  UPON  GROWTH. 


63 


TABLE  V. 

I.  20  per  cent,  of  gas. 

II.  50  per  cent,  of  gas. 

III.  100  per  cent,  of  gas. 

IV.  Control. 

a.  Nitrous  oxide. 

b.  Oxygen. 

Exposed  to  gas  ten  days. 


Is 

o 

3, 

3- 

£ 

e 

B 

d 

_s 

• 

£ 

V 

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t| 

2 

- 

H 

i 

& 

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o 

6 

E 

& 

i 

i 

en 

o 

2 

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2 
bo 

i 

i 

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1 

"o 

"043 
i 

u 

o 

OJ 

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CO   u 

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1  1 

3* 

en 

c* 

•< 

^ 

^ 

"J 

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H 

mm. 

mm. 

111111. 

mm. 

mm. 

days. 

a. 

18 

70 

180.5 

260.5 

590 

44 

2 

2 

•I. 

b. 

10 

60 

210.5 

330 

620 

44 

4 

2 

a. 

7-5 

75 

210.5 

320. 

609 

48 

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I 

II. 

b. 

0 

30 

230. 

44° 

630 

O 

o 

O 

a. 

7 

50 

210.5 

330-5 

625 

38 

3 

1-5 

III. 

b. 

0 

25 

209.5 

315 

615 

44 

4 

2 

29 

85 

250.5 

370-5 

640 

42 

2 

2 

IV. 

control 

18 

65 

150 

280 

580 

43 

3 

i-5 

It  is  to  be  regretted  that  the  root  system  could  not  be  meas- 
ured and  examined  during  the  experiments  without  injuring  the 
plants  to  such  an  extent  as  to  make  the  experiment  useless. 
The  table  does  not  show  anything  in  favor  of  the  plants  treated. 
The  only  conclusion  that  can  be  drawn  is  that  these  gases  have 
no  perceptible  influence  on  Narcissus  bulbs. 

Ammonia. — Vapors  of  this  reagent  are  not  injurious  to  the 
resting  bulb  when  the  amount  of  vapor  present  does  not  exceed 
one  part  in  5000  of  air.  Ariscema  trtphyllum,  Narcissus,  Cro- 
cus, Freesia  and  Tulip  bulbs  were  exposed  to  an  atmosphere 
containing  one  part  of  ammonia  in  5000  of  air  for  ten  days 
without  injuring  the  growing  qualities  of  the  bulb. 


64 


MINNESOTA    BOTANICAL    STUDIES. 


Chloroform. — Vapors  of  chloroform  seem  to  be  very  in- 
jurious to  resting  bulbs.  The  following  quantities  of  the  re- 
agent were  used :  i/iooo,  1/5000  and  i/ioooo  part  in  air  and 
in  all  three  cases  the  bulbs  were  killed.  The  bulbs  used 
were  of  the  same  kind  as  in  the  experiments  with  am- 
monia. The  bulb  decayed  invariably  from  the  shoot  area  to- 
ward the  center  of  the  bulbs  and  never  from  the  root  area. 
The  outer  portion  of  the  bulb  looked  perfectly  natural.  The 
growing  point  of  the  shoot  was  killed  in  every  case.  These 
results  are  of  but  little  value  since  the  temperature  of  the  plant 
house  fell  to  5°  C.  during  one  night. 

Alcohol. — This  reagent  seemed  to  arrest  growth.  Experi- 
ments were  set  up  containing  i/iooo  and  1/500  parts  of  alcohol 
in  4000  cc.  of  air.  The  bulbs  were  kept  under  the  bell  jars  for 
10  days  and  when  taken  out  and  potted  they  were  perfectly 
natural.  The  root  areas  had  begun  to  swell.  No  discolora- 
tion was  noticeable. 

TABLE  VI. 

I.      i/iooo  part  alcohol  in  4000  of  air. 
II.      1/500  part  of  alcohol  in  4000  of  air. 
III.     Control. 


Growth 

Average 

Average 

Average 

Average 

Time  of 

Number 

in  mm. 

growth  in 

growth 

growth 

growth 

first 

of 

when  tak- 

20 days. 

in 

in 

in 

flower. 

flowers. 

en  from 

30  days. 

40  days. 

65  days. 

bell  jar. 

mm. 

mm. 

mm. 

mm. 

I. 

0 

25 

125 

300 

660 

0 

0 

II. 

o 

0 

'3 

20 

3i 

o 

0 

III. 

o 

38 

175 

310 

650 

50 

2 

The  above  table  shows  a  peculiarly  interesting  result.  In  the 
experiment  where  i/iooo  part  of  alcohol  was  used  no  ill  effect 
on  the  plants  could  be  detected.  The  result  is  more  striking  in 
the  experiment  where  1/500  part  of  the  alcohol  was  used.  The 
bulbs  remained  almost  stationary  and  up  to  February  19,  or  100 
days  from  the  time  the  bulbs  were  placed  under  the  bell  jars,  the 
total  growth  was  only  50  mm  Upon  an  examination  it  was  found 
that  the  root  system  was  perfectly  natural  and  well  developed, 
completely  filling  the  four-inch  pots  into  which  the  bulbs  were 


Sandsten :  INFLUENCE  OF  GASES  AND  VAPORS  UPON  GROWTH.    65 

growing.  In  dissecting  the  bulbs  the  floral  structures  were 
found  to  be  very  much  dwarfed  but  the  bud  scales  were  well  de- 
veloped. The  scapes  or  flower  stems  were  greatly  reduced, 
being  only  from  5  to  18  mm.  in  length.  The  bulbs  which  were 
treated  with  i/iooo  part  alcohol  showed  the  same  dwarfed  con- 
dition of  the  floral  organs. 

Bulbs  of  Artscema  triphyllum,  Crocus  and  Tulip  were  treated 
with  the  various  reagents,  but  no  satisfactory  result  was  ob- 
tained. All  the  Crocus  bulbs  died  from  some  unknown  cause 
or  causes.  The  Ariscema  bulbs  were  undoubtedly  affected  by 
the  change  in  temperature  which  occurred  on  the  morning  of 
November  23,  and  to  which  reference  has  previously  been  made. 

5.  THE   INFLUENCE   OF   GASES    UPON    GROWTH   OF   PLANTS 
GROWN  IN  NUTRIENT  SOLUTION  OF  WATER  CULTURE. 

A.  Land  plants  grown  in  nutrient  solution.     Seedlings  of 
Zea  mats  were  used  in  these  experiments  and  the  method  de- 
scribed in  the  introduction  was  observed.     The  duration  of  the 
experiment  was  limited  to  eighteen  days.     The  average  growth 
for  this  period,  taken  in  four  separate  experiments  in  which  the 
seedlings  were  grown  in  a  nutrient  solution  saturated  with  com- 
mercial nitrous  oxide,  was  203  mm.  for  the  roots  and  209  for 
the  shoots.     The  control  plants  grown  in  nutrient  solution  with- 
out nitrous  oxide  showed  an  average  growth  of  213  mm.  for 
the  roots  and  165  mm.  for  the  shoots.     The  result  shows  a  gain 
in  favor  of  the  nitrous  oxide  in  the  shoot  and  a  loss  for  the  root, 
but  the  result  needs  verification. 

B.  Aquatic  -plants  grown   in   river  water.     Wide  bell  jars 
were  inverted  and  filled  with  about  one  inch  of  soil  over  which 
a  thin  layer  of  clean  sand  was  spread  ;  in  this  substratum  several 
plants  of  Philotria  were  planted  and  the  bell  jar  filled  about 
half  full  of  river  water.     The  water  in  the  bell  jars  was  kept 
saturated  with  nitrous  oxide  by  means  of  inverted  bottles  which 
were  first  filled  and  inserted  under  water  and  this  water  was  dis- 
placed by  nitrous  oxide.     The  bottles  were  kept  in  position  by 
means  of  iron  stands.     The  bottles  were  refilled  as  soon  as  the 
gas  was  exhausted. 


66 


MINNESOTA    BOTANICAL    STUDIES. 


TABLE  VII. 

I.    Philotria  in  a  saturated  solution  of  N2O. 
II.   Control. 


Average  growth 
in  5  days, 
mm. 

Average  growth 
in  10  days, 
mm. 

Average  growth 
in  20  days, 
mm. 

Average  growth 
in  30  days, 
mm. 

I.  • 

8 

25 

53 

68 

II. 

3 

9 

18.5 

29 

The  marked  results  shown  in  the  above  table  were  duplicated 
in  all  the  experiments  set  up.  The  average  growth  of  the 
Philotria  branches  after  28  days  taken  in  another  test  from 
four  experiments  was  17  mm.  and  the  average  growth  from 
the  control  was  11.5  mm.;  showing  conclusively  that  nitrous 
oxide  has  a  stimulating  effect  on  Philotria. 

To  test  further  the  effect  of  nitrous  oxide  on  aquatic  plants, 
stems  of  Salvinia  natans,  having  an  aggregate  of  80  leaves 
were  placed  in  bell  jars  set  up  in  the  same  manner  as  in  the  pre- 
vious experiment  and  the  water  kept  saturated  with  gas.  The 
experiments  were  allowed  to  run  for  40  days,  when  the  number 
of  leaves  were  counted.  The  control  or  checks  were  set  up 
in  exactly  the  same  manner  as  the  tests  with  the  exception  of 
nitrous  oxide. 

TABLE  VIII. 

I.     Salvinia  in  a  saturated  solution  of  NaO. 
II.     Control. 


Number  of  leaves 
at  the  beginning 
of  the  experiment. 

Number  of  leaves 
at  the  close  of 
the  experiment. 

Total  gain  in 
number  of 
leaves  in  40  days. 

a 

80 

131 

5I 

a 
I.        a 
a 

80 
80 
80 

118 
114 
1  08 

38 

34 

28 

TT        a 
a 

80 
80 

99 
104 

19 
24 

The   result  of   every  experiment    showed  that    growth    was 
accelerated  by  nitrous  oxide. 


Sandsten:  INFLUENCE  OF  GASES  AND  VAPORS  UPON  GROWTH.     67 

Ammonia  was  tried  on  the  same  water  plants  under  exactly  the 
same  conditions  as  above.  One-tenth  of  a  cc.  in  2000  cc.  of 
water,  and  one-five-hundredth  of  a  cc.  in  2000  cc.  of  water 
were  used,  but  in  both  cases  both  the  Sahnnia  and  Philotrii 
plants  died. 

CONCLUSIONS. 

Influence  of  Gases. 

From  the  foregoing  tables  and  records  the  following  conclu- 
sions seem  to  be  warranted  : 

Nitrous  oxide.  Seeds  of  Phaseolus  multiflorus  and  Vicia 
faba  will  not  germinate  in  an  atmosphere  containing  80%  of 
nitrous  oxide.  Seedling  of  Phaseolus  multiflorus  and  Vicia  faba 
will  remain  active  more  than  24  hours  in  an  atmosphere  of  com- 
mercial nitrous  oxide,  but  no  growth  can  take  place.  Shoots 
exhibit  accelerated  growth  after  being  kept  in  an  atmosphere  of 
free  N?O  or  in  an  atmosphere  where  the  amount  of  gas  ranges 
from  25  to  100  per  cent.  No  growth  in  shoots  could  be  detected 
during  the  confinement  under  the  bell  jars. 

Water  plants  such  as  Salvinia  natans  and  Philotria  show  in- 
creased growth  in  solutions  saturated  with  N2O. 

Oxygen.  Seeds  readily  germinate  in  an  atmosphere  of  free 
oxygen.  Seedlings  kept  in  an  atmosphere  of  free  oxygen  do 
not  grow  as  rapidly  as  seedlings  in  a  moist  chamber  containing 
ordinary  air.  Growing  shoots  kept  in  an  atmosphere  contain- 
ing from  25  to  100  per  cent,  of  free  oxygen  will  remain  unal- 
tered as  long  as  20  days,  but  on  removal  slowly  perish. 

Influence   of  Vapors. 

Ammonia  (NOH4). — Vapors  of  this  reagent  when  used  in  quan- 
tities not  exceeding  1/24000  part  are  not  harmful  to  the  germin- 
tion  of  seeds  of  Phaseolus  multiflorus.  Seeds  exposed  for  nine 
days  in  glass  chambers  containing  from  1/24000  to  1/32000  parts 
of  NOH4  germinated  as  freely  as  the  control.  The  seed  of 
Vicia  faba  is  very  susceptible  to  the  influence  of  this  reagent 
and  seeds  kept  in  a  glass  chamber  for  nine  days  containing 
1/28000  part  of  NOH4  failed  to  germinate.  In  an  atmosphere 
containing  1/32000  part  of  NOH4  90  per  cent,  of  the  seed  ger- 
minated. 

Seeds  of   Phaseolus  multiflorus  and    Vicia  faba  kept  for  nine 


68  MINNESOTA    BOTANICAL    STUDIES. 

days  in  an  atmosphere  containing  1/20000  of  NOH4  failed  to  ger- 
minate. The  growth  of  young  seedlings  of  Zea  mats  kept  in 
a  moist  chamber  for  48  hours  containing  1/20000  partNOH4  was 
retarded.  Growing  shoots  are  badly  affected  when  kept  in  an 
atmosphere  containing  1/15000  part  of  NOH4.  Resting  bulbs 
are  not  effected  by  being  kept  in  an  atmosphere  containing  one 
part  of  NOH4  in  5000  of  air.  Salvinia  natans  and  Philotria  are 
killed  by  introducing  .1  cc.  of  NOH4  to  every  2000  cc.  of 
water. 

Chloroform  and  Ether. — These  two  reagents  have  a  very 
similar  effect  upon  growth.  Seedlings  of  Zea  mats  kept  in  a 
moist  chamber  containing  I/TOOOO  part  of  chloroform  or  ether 
show  a  marked  acceleration  in  growth  after  release.  In  an  at- 
mosphere containing  1/5000  growth  is  greatly  retarded.  Rest- 
ing bulbs  and  growing  shoots  are  equally  susceptible  and  are 
killed  after  being  exposed  for  ten  to  twenty  days  in  an  atmos- 
phere containing  i/ioooo  part  of  the  reagent. 

Carbon  bisulphide. — The  smallest  trace  of  carbon  bisulphide 
present  is  injurious  to  growing  plants,  although,  as  G.  Hicks 
(III.)  has  shown,  it  is  inoperative  on  resting  seeds. 

Alcohol  has  no  effect  upon  the  growth  of  seedlings  when 
used  in  quantities  not  exceeding  i/ioooo.  If  the  larger  quanti- 
ties are  used  the  growth  is  retarded  and  the  seedlings  are  killed. 
Resting  bulbs  kept  in  an  atmosphere  containing  i/ioooto  1/500 
parts  of  alcohol  grew,  but  the  floral  organs  were  dwarfed  and 
the  buds  remained  unopened. 

LITERATURE  TO  WHICH  REFERENCE  is  MADE. 

I.  Detmer.  Ueber  die  Einwirkung  verschiedener  Gase  insbe- 
sondere  des  Stickstoffoxydulgases  auf  Pflanzen.  Landw.  Jahrb. —  : 
213.  1882. 

II.  Detmer.     Das  Verhalten  der  Pflanzen  im  Contact   mit  Stick- 
stoffoxydulgase.     Physiol.     Prakt.     2  Ed.,  p.  235.      1895. 

III.  Hicks.     Bull— Bot.  Div.     Dept.  of  Agric.— 1896. 

IV.  Johannsen — Bot.   Cent. — 1896. 

V.  Moeller.       Ueber    Pflanzenathmung.      I    Das  Verhalten    der 
Pflanzen  zu  Stickstoffoxydul.     Ber.  d.  deut.  hot.  Ges.  2:  35.      1884. 
VI.  Townsend  C.  O.     Correlation  of  growth   under  the  influence 
of  injuries.      Annals  of  Botany,      n:     509.      1897. 


VIII.    SEEDLINGS    OF   CERTAIN   WOODY   PLANTS. 


FRANCIS  RAMALEY. 


The  following  observations  on  seedlings  of  woody  plants 
were  made  at  the  University  of  Minnesota,  during  the  years  1896, 
1897  and  1898.  The  plants  were  grown  from  seed  either  col- 
lected by  the  writer,  or  obtained  from  reliable  dealers. 

Most  of  the  species  studied  have  not  hitherto  been  investi- 
gated. The  author  has,  however,  re-examined  some  plants  de- 
scribed by  former  investigators,  especially  in  cases  where  the 
printed  descriptions  were  incomplete  or  without  illustrations. 

The  measurements  given  are  in  all  cases  based  on  a  consider- 
able number  of  plants  examined.  It  has  been  found  that  the 
exact  lower  limit  of  the  hypocotyl  is  not  always  'readily  deter- 
mined, although,  generally  it  is  enough  larger  in  diameter  than 
the  root  to  be  exactly  located.  It  has  seemed  best  in  giving  the 
length  of  the  hypocotyl  to  measure  its  full  extent  rather  than 
simply  that  part  above  the  ground. 

An  attempt  has  been  made  to  note,  as  far  as  possible,  whether 
the  seed  coat  is  carried  up  or  remains  underground  and  also 
how  much  the  cotyledons  increase  in  size  after  appearing  above 
the  ground.  These  points  have  not  generally  been  noted  by 
students  of  seedlings. 

In  most  cases  the  length  of  time  required  for  germination  of 
the  seeds  is  given.  The  figures  are  for  the  first  seedlings. 
Oftentimes  plants  will  appear  every  few  days  for  over  a  month 
after  the  first  have  come  up.  Unless  otherwise  stated  it  is  to  be 
understood  that  the  seeds  were  planted  in  the  following  spring 
after  ripening.  The  plants  studied  will  be  considered  in  the 
order  of  Engler  and  Prantl. 

SALICACE.E. 
Populus  deltoides  MARSH. 

The  seed  of  the  "  cottonwood  "  ripens  in  June  and  should  be 
planted  at  once.  The  young  plants  appear  above  ground  in  a 
week  or  sooner. 


70  MINNESOTA    BOTANICAL    STUDIES. 

The  cotyledons  are  petiolate,  the  blade  being  ovate-oblong, 
about  5  mm.  in  length  and  4  mm.  broad.  The  petiole  is  about 
3  mm.  long.  There  is  but  little  increase  in  size  as  the  plant 
grows  older. 

The  first  two  leaves  are  opposite,  lanceolate,  short-petiolate, 
of  willow-like  shape.  They  are  about  10  mm.  long  before  the 
epicotyl  has  developed  at  all  and  do  not  afterward  increase  in 
size.  The  hypocotyl  is  10-15  mm.  long;  the  epicotyl  reaches 
a  length  of  8  or  10  mm. 

The  third  and  fourth  leaves  are  nearly  opposite ;  later  ones 
are  alternate.  The  later  leaves  become  broader  and  longer 
petioled,  gradually  assuming  the  deltoid  form  characteristic  of 
the  species. 

ULMACE^. 
Ulmus  americana  LINN. 

The  "  white  elm  "  is  a  native  of  the  eastern  and  central  United 
States.  The  seeds  ripen  in  early  spring  and  must  be  sown  at 
once.  They  germinate  in  about  a  week.  Often,  but  not  al- 
ways, the  pericarp  is  carried  up  by  the  growing  seedling. 

The  cotyledons  are  at  first  obovate,  slightly  auriculate,  5  mm. 
long  and  2-3  mm.  broad.  They  are  indistinctly  reticulately 
veined.  They  increase  but  slightly  in  size  and  seldom  become 
more  than  7  mm.  long.  They  are  short-petioled.  The  hypo- 
cotyl is  slender,  not  enlarged  at  the  base,  25-35  mm.  long, 
but  in  time  it  may  reach  a  length  of  50  mm.  The  epicotyl  is 
about  10  mm.  in  length. 

The  leaves  are  petiolate,  ovate,  coarsely  serrate,  with  distinct 
veining.  The  first  two  are  opposite,  the  third  and  fourth  nearly 
so.  Later  leaves  are  alternate. 

Ulmus  fulva  MICHX. 

The  "slippery  elm,"  like  the  previously  described  species, 
ripens  its  seeds  early  in  the  spring.  These,  when  planted  at 
once,  germinate  in  about  two  weeks.  The  seedling  resembles 
that  of  U.  Americana  in  all  essential  respects. 

Celtis  occidentalis  LINN. 

This  is  a  fine  tree  native  to  the  central  United  States  and 
Canada.  It  is  known  as  the  "  hackberry."  The  seeds  germi- 
nate in  from  four  to  six  weeks.  The  seed  coat  remains  under 
ground. 


Ramaley :     SEEDLINGS  OF  CERTAIN  WOODY  PLANTS.  71 

The  cotyledons  are  at  first  10  mm.  long,  but  by  the  time  the 
first  pair  of  foliage  leaves  appear  they  are  30  mm.  in  length, 
17  mm.  in  width,  ovate,  entire,  notched  at  the  apex.  The 
epicotyl  is  10  mm.  in  length.  The  figures  here  given  are  sub- 
stantially correct  for  all  the  plants  examined  by  the  present 
writer.  Lubbock's  *  figures  for  seedlings  of  this  species  are 
about  one-half  those  here  given. 

The  epicotyl  is  at  length  20  mm.  long.  The  first  two  foliage 
leaves  are  opposite,  the  later  ones  alternate.  Leaves  of  the  first 
year  are  not  at  all  conspicuously  oblique  at  base  as  are  those  of 
older  plants. 

MORACE^E. 
Toxylon  pomiferum  RAF. 

This  is  the  well-known  "  osage  orange"  of  the  south-central 
United  States.  The  seeds  germinate  in  about  one  month  after 
planting.  The  seed  coat  is  often  carried  up  by  the  cotyledons 
which  are  thus  prevented  from  opening  till  they  have  increased 
somewhat  in  size. 

When  they  first  appear  the  cotyledons  are  9  mm.  long  and  5 
mm.  broad,  obovate-oblong,  entire,  short-petioled.  The  hypo- 
cotyl  is  stout,  35-50  mm.  long. 

The  cotyledons  grow  rapidly  in  size  and  by  the  time  the  first 
leaves  are  well  developed  have  increased  to  20  mm.  in  length 
and  12  mm.  in  width.  The  petiole  is  distinctly  margined  and 
4  mm.  long.  The  veining  of  the  blade  is  distinct.  The  epi- 
cotyl is  10-15  mm.  long.  The  first  two  foliage  leaves  are  op- 
posite, narrowly  lanceolate,  ovate,  entire,  or  nearly  so,  distinctly 
veined.  The  later  leaves  are  alternate,  often  pointed  at  the 
base  as  well  as  the  apex.  The  seedling  of  this  plant  was 
studied  by  Lubbock,f  but  not  figured. 

Broussonetia  papyrifera  (LINN.)  VENT. 

The  seeds  of  this  oriental  tree,  the  "  paper  mulberry,"  ger- 
minate in  about  three  or  four  weeks  after  planting.  The  seed 
coat  is  carried  up  and  often  remains  attached  to  one  of  the 
cotyledons  for  a  time  after  they  have  opened. 

The    hypocotyl  is    rather    slender,   12-15  mm.   long.     The 

•On  Seedlings,  2:  493.     1892. 
t  Op.  cit.  a  :  498. 


72  MINNESOTA    BOTANICAL    STUDIES. 

cotyledons  are  oval ;  when  fully  open  they  have  a  petiole  2  mm. 
long  and  blade  8  mm.  long  and  5  mm.  broad,  very  slightly 
notched  at  the  apex.  When  they  first  emerge  from  the  seed 
coat  the  leaves  are  not  over  5  mm.  in  length. 

The  first  foliage  leaves  are  opposite  ;  they  are  petiolate,  nar- 
rowly ovate,  serrate,  slightly  heart-shaped  at  base  and  more 
nearly  entire  than  the  later  leaves  which  are  alternate,  long- 
petioled,  serrate  and  frequently  more  or  less  two-  or  three- 
lobed  or  parted.  Usually  about  the  close  of  the  second  sea- 
son the  well-known  peculiar  characteristic  leaves  make  their 
appearance. 

MAGNOLIACE^E. 
Liriodendron  tulipifera  LINN. 

The  seeds  of  the  "  tulip  tree  "  germinate  in  from  four  to  six 
weeks  after  planting.  The  wing-like  pericarp  remains  in  the 
soil. 

The  cotyledons  when  they  first  appear  are  about  7  mm.  long 
and  5  mm.  broad,  almost  sessile,  ovate-oblong  in  shape.  Before 
the  first  leaf  appears  each  cotyledon  has  developed  a  distinct 
petiole  2  or  3  mm.  long,  while  the  blade  is  about  12  mm.  in 
length. 

The  foliage  leaves  are  alternate.  The  first  is  broadly  ovate- 
oblong,  petiolate,  emarginate,  with  entire  margin.  The  second 
and  third  resemble  the  first.  The  characteristic  leaves  appear 
toward  the  close  of  the  first  season  or  not  till  the  second  year. 
The  epicotyl  is  extremely  short,  1—2  mm.  long.  Succeeding 
internodes  are  likewise  short. 

CALYCANTHACE^E. 
Butneria  florida  (LINN.)  KEARNEY. 

This  is  the  familiar  "sweet-scented  shrub"  commonly  culti- 
vated in  the  eastern  United  States.  It  is  native  from  Virginia 
to  the  Gulf  of  Mexico.  The  seeds  require  a  month  or  more  to 
germinate.  The  cotyledons  are  rolled  longitudinally  about 
each  other  in  the  seed  and  remain  rolled  up  for  two  or  three 
days  after  appearing  above  ground. 

The  hypocotyl  is  stout,  20  mm.  long.  Cotyledons  are  thick, 
dark  green,  slightly  auriculate  at  base,  petiolate,  generally 
somewhat  trapezoidal,  the  apex  broadly  incurved.  They  are  at 


Ramaley  :     SEEDLINGS  OF  CERTAIN  WOODY  PLANTS.          73 

first  about  12-15  mm-  l°ng  and  25  rnm.  broad.  Eventuall}' 
they  may  become  20  mm.  long  and  30  mm.  broad  with  petioles 
10  mm.  in  length.  In  shape  they  are  often  quite  asymmetrical. 
The  foliage  leaves  are  opposite,  ovate,  pointed,  entire.  The 
first  do  not  differ  materially  from  the  later  ones.  The  epicotyl 
is  about  the  same  length  as  the  hypocotyl. 

Butneria  fertilis  (WALT.)  KEARNEY. 

The  seedling  of  this  plant  does  not  differ  in  any  important 
respect  from  that  of  the  species  just  described. 

CAESALPINACE^E. 

Parkinsonia  aculeata  LINN. 

The  seeds  of  this  shrub  germinate  in  about  two  weeks  after 
planting.  The  seed  coat  is  usually  carried  up. 

When  they  first  appear  the  cotyledons  are  15  mm.  long,  8 
mm.  broad,  ovate-elliptical,  sessile,  very  slightly  auriculate  at 
base.  The  hypocotyl  is  stout,  30-50  mm.  in  length.  The  coty- 
ledons increase  in  size  until  they  are  25  mm.  long. 

Foliage  leaves  are  alternate  ;  all  are  pinnate,  the  first  has  five 
pairs,  the  second  six  pairs  of  leaflets.  The  epicotyl  is  9  mm. 
long  when  two  leaves  have  appeared. 

Cercis  canadensis  LINN. 

This  is  the  well-known  "  red  bud  "  or  "Judas  tree  "  of  the 
central  United  States.  The  seeds  germinate  in  about  two 
weeks.  The  seed  coat  is  usually  carried  up,  holding  the  coty- 
ledons together  until  erect.  The  veins  of  the  cotyledons  are 
distinct  even  before  the  cotyledons  have  separated. 

The  cotyledons  are  broadly  ovate,  at  first  6  mm.  long  and  4 
mm.  broad,  eventually  15  mm.  long  and  8  mm.  broad.  The 
hypocotyl  is  stout,  10-30  mm.  long.  This  is  of  interest  since 
the  hypocotyl  of  C.  siliquastrum  WILLD.,  as  described  by  Lub- 
bock,*  is  but  5-6  mm.  in  length. 

The  epicotyl  is  20-30  mm.  long.  Foliage  leaves  are  all  al- 
ternate, entire,  cordate,  long-petioled. 

Gleditsia  triacanthos  LINN. 

The  "  honey  locust,"  as  this  plant  is  called,  is  a  familiar  tree 
of  the  central  United  States.  The  seeds  germinate  in  about  one 

*  Op.  cit.  i :  465. 


74  MINNESOTA    BOTANICAL    STUDIES. 

month  after  sowing.  The  seed  coat  is  sometimes  carried  above 
ground,  but  it  as  often  remains  in  the  soil. 

The  hypocotyl  is  stout,  25-30  mm.  in  length.  The  cotyle- 
dons are  sessile,  slightly  auriculate,  oblong,  18  mm.  in  length 
and  9  mm.  broad.  They  do  not  increase  greatly  in  size. 

Leaves  are  alternate  and  pinnate.  The  second  appears  be- 
fore the  first  is  fully  open.  The  first  leaf  usually  has  eight  pairs 
of  leaflets,  the  second  has  eleven  pairs,  the  third  thirteen  pairs. 
When  these  leaves  have  developed  the  hypocotyl  is  about  50 
mm.  long,  the  epicotyl  20-25  mm. 

The  first  leaves  are  described  by  Tubeuf  *  as  having  ten  pairs 
of  leaflets.  In  the  plants  examined  by  the  present  writer  the 
first  leaf  had  never  more  than  nine  pairs  of  leaflets. 

PAPILIONACE^. 
Amorpha  fruticosa  LINN. 

This  is  an  ornamental  shrub  indigenous  to  North  America 
and  frequently  cultivated.  The  seeds  germinate  in  about  two 
weeks  after  planting. 

When  the  cotyledons  first  appear  they  are  ovate  in  shape, 
about  5  mm.  long  and  2.5  mm.  broad.  By  the  time  they  are 
fully  open  they  measure  8  mm.  in  length.  The  hypocotyl  at 
this  time  is  25  mm.  long,  quite  slender,  gradually  thickened 
below. 

The  cotyledons  attain  a  length  of  12  mm.  They  are  sessile. 
The  epicotyl  is  15  mm.  in  length.  Foliage  leaves  are  alternate. 
The  first  five  or  six  are  simple,  broadly  ovate,  petiolate.  After 
these  the  leaves  are,  for  a  space,  pinnately  trifoliate.  The 
terminal  leaflet  is  larger  and  longer  stalked  than  the  lateral  ones. 
Later  leaves  are  pinnate  with  numerous  leaflets. 

Amorpha  nana  NUTT. 

The  seedlings  of  this  shrub  resemble  those  of  A.  fruticosa  save 
that  they  are  much  smaller.  The  hypocotyl  does  not  become 
more  than  8  or  10  mm.  in  length  and  the  epicotyl  is  only  about 
5  mm.  long.  Cotyledons  are  5  mm.  long  and  3  mm.  broad. 

Leaves  are  alternate.  The  first  six  to  ten  are  simple.  They 
are  broadly  orbicular,  emarginate,  petiolate,  with  a  distinct  mid- 
vein.  As  in  the  former  species  the  later  leaves  are  pinnately 
compound. 

*  Samen,  Fruchte  und  Keimlinge,  12".    1891. 


Ramaley :     SEEDLINGS  OF  CERTAIN  WOODY  PLANTS.          76 

The  writer  is  indebted  to  Mr.  D.  M.  Andrews,  of  Boulder, 
Colo.,  for  seedlings  of  this  plant  and  of  Acer  glabr ion. 

Robinia  pseudacacia  LINN. 

The  "  locust  tree"  is  a  native  of  the  middle  and  southeastern 
United  States.  The  seeds  germinate  in  about  two  weeks  after 
planting. 

Seedlings  of  this  plant  were  studied  by  Lubbock  *  and  by 
Flot  f  but  the  fact  is  not  stated  by  these  writers  that  the  coty- 
ledons are  at  first  somewhat  narrowly  elliptical  or  obovate  and 
only  at  a  rather  late  stage  become  "  oblong-oval."  The  descrip- 
tions hitherto  published  have  not  been  accompanied  by  satis- 
factory figures. 

RUTACE^E. 
Ptelea  trifoliata  LINN. 

This  is  the  so-called  "  hop  tree"  of  the  central  United  States. 
The  seeds  germinate  in  from  three  to  four  weeks,  the  seed  coat 
remaining  underground.  Almost  as  soon  as  the  cotyledons  get 
above  ground  they  become  erect  and  then  separate. 

The  cotyledons  are  nearly  sessile,  elliptical-oblong,  entire, 
6  mm.  long  and  3  mm.  broad.  They  grow  for  some  time  and 
become  at  length  18  mm.  long,  7  mm.  broad,  minutely  serrate, 
short-petiolate,  with  midvein  distinct.  The  hypocot}*!  is  rather 
stout,  15-20  mm.  long. 

The  epicotyl  is  20-40  mm.  long  when  the  first  leaf  is  fully 
open.  It  often  elongates  somewhat  after  that  time.  Leaves 
are  alternate.  The  first  foliage  leaf  is  usually  simple,  ovate, 
petiolate,  with  crenulate  margin.  Sometimes  it  is  trifoliate, 
sometimes  but  partially  compound ;  perhaps  one  of  the  side 
leaflets  is  separate,  but  not  the  other.  The  second  leaf  is 
usually  trifoliate,  sometimes  incompletely  so.  Later  leaves  are 
all  trifoliate,  the  terminal  leaflet  larger  than  the  lateral  ones. 

A  description  of  this  seedling  is  given  by  Lubbock  $  but  there 
is  no  figure. 


*  Op.  cit.,  i  :  422. 

tFlot.     Recherches  sur  la  structure  comparee  de  la  tige  des  arbres. — Rev. 
gen.  de  Bot.  2  :  20.     1890. 
JOp.  cit.  I  :  322. 


76  MINNESOTA    BOTANICAL    STUDIES. 

SIMARUBACE.E. 
Ailanthus  glandulosa  DESF. 

This  well-known  tree  is  a  native  of  eastern  Asia.  It  is,  how- 
ever, grown  extensively  in  this  country.  The  writer's  observa- 
tions on  the  seedlings  differ  somewhat  from  those  previously 
published.* 

The  seeds,  planted  in  May,  germinated  in  from  two  to  three 
weeks.  The  seed  coat  and  wing  sometimes  remain  in  the 
ground  but  are  quite  often  carried  up  by  the  elongation  of  the 
hypocotyl  before  the  cotyledons  emerge. 

The  cotyledons  are  at  first  about  6  mm.  long.  By  the  time 
they  are  fully  open  they  have  increased  somewhat  in  size  and 
the  hypocotyl  has  attained  its  full  length,  viz.  about  40  mm. 
When  the  first  leaves  have  opened  the  cotyledons  are  broadly 
obovate,  petiolate,  with  the  blade  15  mm.  long,  n  mm.  broad 
and  the  petiole  5  mm.  long. 

The  epicotyl  is  at  length  20  mm.  long.  The  first  few  leaves 
are  trifoliate.  Later  leaves  are  pinnate. 

ANACARDIACE^:. 
Schinus  molle  LINN. 

This  is  the  so-called  "pepper-tree"  sometimes  planted  in 
California  but  a  native  of  tropical  America.  A  description  of 
the  germination  of  the  seed  is  given  by  Lubbock,f  whose  ac- 
count in  this  case,  the  present  writer  only  desires  to  supplement. 

The  cotyledons  are  remarkable  for  their  great  increase  in 
size ;  beginning  with  a  length  of  5  mm.  the  blade  is  finally  20 
-25  mm.  long  and  broad  in  proportion.  The  petiole  is  about 
4  mm.  in  length. 

CELASTRACE^E. 
Celastrus  scandens  LINN. 

This  is  the  "  climbing  bitter-sweet, T  a  common  native  liana 
of  the  United  States.  It  is  frequently  cultivated.  The  seeds 
ripen  in  the  fall.  If  planted  the  following  spring  they  usually 
require  a  year  to  germinate. 

The  cotyledons  are  thin,  reticulately  veined,  petiolate,  oval- 
oblong  in  shape.  At  first  they  are  10  mm.  long  and  5  mm. 

*Lubbock  op.  cit.  i  :  327. 
|Op.  cit.  2:  335. 


Ramaley :     SEEDLINGS  OF  CERTAIN  WOODY  PLANTS.  77 

broad,  but  grow  rapidly  and  become  about  twice  that  size.  As 
they  grow  older  they  become  broader  in  proportion  to  the  length. 
The  petiole  is  finally  about  5  mm.  long.  The  hypocotyl  tapers 
gradually  to  the  root  so  that  its  exact  limit  is  not  easily  recog- 
nized. It  reaches  a  length  of  40  or  50  mm. 

The  epicotyl  is  about  15  mm.  long.  Leaves  are  all  alternate. 
The  first  leaves  are  not  different  from  those  formed  later. 

ACERACE^E. 
Acer  negundo  LINN. 

Seeds  of  the  "box-elder"  germinate  in  from  one  to  two 
weeks  after  sowing.  The  large  winged  pericarp  is  brought 
above  ground. 

The  hypocotyl  is  25-35  mm.  long  when  the  cotyledons  first 
open  and  does  not  grow  longer.  The  cotyledons  are  strap- 
shaped,  sessile,  entire,  tri-veined  ;  about  30  mm.  long  and  5  mm. 
broad. 

The  epicotyl  becomes  5-8  mm.  long.  Leaves  are  opposite. 
The  first  two  are  ovate,  acute,  serrate,  petiolate.  Later  ones 
are  tri-cleft.  Usually  the  sixth  or  seventh  pair  and  all  later 
ones  are  pinnately  compound. 

Acer  glabrum  TORR. 

The  seedling  of  the  "  Rocky  mountain  maple  "  resembles  that 
of  A.  negundo.  The  hypocotyl  is  shorter,  20  mm.  long,  and 
the  cotyledons  about  20  mm.  long,  5  mm.  broad.  Leaves  are 
opposite,  long-petioled,  ovate-cordate,  the  second  pair  somewhat 
lobed.  Later  leaves  are  three-  or  five-lobed,  the  lobes  more  or 
less  acute  and  sharply  serrate. 

Acer  saccharinum  LINN. 

The  seeds  of  the  "  soft  maple"  germinate  in  about  ten  days 
after  planting.  The  first  leaves  are  well  developed  when  the 
plant  appears  above  ground.  The  cotyledons  remain  in  the 
soil  for  a  time  enclosed  in  the  pericarp  which  eventually  decays. 
Sometimes  they  do  not  appear  above  ground  *  at  all. 

The  hypocotyl  is  stout,  about  20  mm.  long.  The  cotyledons 
are  somewhat  fleshy,  asymmetrical,  short-petioled,  bent  around 

*Winkler.  Kleinere  morph.  Mittheilungen,  in  Verhandl.  d.  Bot.  Ver.  d. 
Provinz  Brandenburg,  18  :  99.  1877. 


78  MINNESOTA    BOTANICAL    STUDIES. 

so  that  both  are  on  the  same  side  of  the  stem.     They  are  about 
16  mm.  long  and  8  mm.  broad. 

The  epicotyl  often  becomes  greatly  elongated,  reaching  a 
length  of  50-100  mm.  Leaves  are  all  opposite,  those  of  the 
seedling  are  the  same  shape  as  the  later  leaves. 

RHAMNACE.E. 
Berchemia  racemosa  SIEB.  &  Zucc. 

This  plant  is  a  shrub  with  conspicuously  veined  leaves.  It 
is  a  native  of  Asia.  The  seeds  require  two  or  three  weeks  to 
germinate.  It  is  often  a  number  of  days  before  the  cotyledons 
get  out  of  the  seed  coat  which  is  carried  above  ground. 

When  the  cotyledons  first  emerge  they  are  sessile,  strap- 
shaped,  8  mm.  long  and  i^  mm.  broad.  They  remain  about  the 
same  size  for  a  time  after  they  are  fully  open.  The  hypocotyl 
is  slender,  about  15  mm.  long. 

By  the  time  the  first  foliage  leaves  are  open  the  hypocotyl  is 
15-20  mm.  long  ;  the  epicotyl  5  mm.  in  length  and  the  cotyledons 
10  or  12  mm.  long  and  1.5-2  mm.  broad.  The  foliage  leaves 
are  ovate,  petiolate,  conspicuously  veined  ;  the  first  two  are 
opposite  or  nearly  so,  all  others  are  alternate. 

Rhamnus  purshiana  DC. 

This  is  a  handsome  tree  of  Pacific  North  America  sometimes 
planted  in  the  eastern  United  States.  The  bark  is  the  "  Cas- 
cara  Sagrada  "  of  the  drug  stores.  The  seeds  require  a  month 
or  more  to  germinate. 

The  cotyledons  increase  but  slightly  in  size  after  opening. 
They  are  obovate,  entire,  sessile  or  nearly  so,  7  mm.  long  and 
5  mm.  broad.  The  hypocotyl  is  25-30  mm.  long. 

The  epicotyl  is  slender,  15—20  mm.  long.  Foliage  leaves  are 
ovate,  pointed,  petiolate,  alternate.  The  first  two  and  the  third 
and  fourth  are,  however,  nearly  opposite.  The  margin  of  the 
leaf  is  finely  serrate  ;  the  veining  very  prominent. 


Vitis  cordifolia  MICHX. 

This  is  one  of  the  commonest  wild  grapes  found  in  the  north- 
ern United  States.     The  seeds  germinate  readily,  the   cotyle- 


Ramaley:     SEEDLINGS  OF  CEKTAIX  WOODY  PLAVTS.          79 

dons  appearing  above  ground  in  about  four  weeks.  Sometimes 
the  seeds  do  not  germinate  till  the  second  year. 

The  cotyledons  are  ovate,  petiolate,  veined.  When  they  first 
appear  the  blades  are  about  10  mm.  long  and  6  mm.  broad. 
They  grow  to  about  18  mm.  in  length,  and  a  corresponding  width 
before  the  first  leaves  appear,  after  which  time  they  do  not  in- 
crease in  size.  The  petiole  is  about  8  mm.  long.  The  hypo- 
•  •".  is  stout,  from  25—30  mm.  long ;  it  does  not  grow  longer. 

The  leaves  are  all  alternate,  ovate-heart-shaped,  irregularly 
dentate,  palmately  5-veined.  When  the  first  leaf  appears  the 
epicotvl  is  about  8  mm.  long.  It  may  eventually  reach  a  length 
of  10  or  12  mm. 

Parthenocisstts  quinquefolia  (Lrxx.)  PLANCH. 

This  is  the  familiar  **  Virginia  creeper,"  a  native  of  the 
United  States  and  frequently  planted.  Seeds  germinate  in 
about  three  wee 

.e  hvpocotyl  is  stout,  from  20-40  mm.  long.  The  cotyle- 
dons are  long-petiolate.  The  blade  is  cordate,  prominently 
veined,  at  length  20  mm.  long,  20  mm.  broad.  The  petiole  is 
channeled,  20  mm.  long.  Both  hypocotyl  and  petioles  are  pink 
except  that  part  of  the  hvpocotyl  which  is  below  ground.  The 
hvpocotyl  becomes  verv  much  thickened  toward  the  end  of  the 
season,  exhibiting  a  well-marked  *'  region  tigellaire." 

The  epicotyl  is  undeveloped,  the  first  leaf  arising  just  above 
the  cotyledons.  Leaves  are  all  alternate  and  quinquefoliate 
from  the  beginning.  The  first  do  not  differ  from  the  later 
ones. 

STERCULIACRE. 

Sterculia  platanifolia.  Lixx. 

The  seeds  of  this  oriental  tree  germinate  in  about  a  month 
after  planting.  A  part  of  the  seed  coat  is  often  attached  to  the 
cotyledons  when  they  first  appear  above  the  ground. 

The  hypocotyl  is  stout,  40  mm.  long  at  the  time  the  cotyle- 
dons open.  These  are  broadly  oblong  or  orbicular,  entire, 
slightly  cordate  at  base,  with  petioles  nearly  as  long  as  the 
blades.  The  latter  are  at  first  about  18  mm.  long  and  16  mm. 
broad  but  become  very  large,  sometimes  40  mm.  long  and  45  mm. 
broad.  They  are  palmately  five-veined.  The  midvein  forks 
some  distance  from  the  apex. 


80  MINNESOTA    BOTANICAL    STUDIES. 

The  epicotyl  is  about  10  mm.  long.  Leaves  are  alternate. 
The  first  leaf  is  broadly  heart-shaped,  entire,  petiolate,  palmately 
five-veined ;  the  midvein  runs  to  the  apex  of  the  leaf. 

EL^EAGNACE^E. 

Elaeagnus  umbellata  THUNB. 

The  seeds  of  this  Japanese  shrub  require  about  four  weeks  to 
germinate.  The  seed  coat  is  often  carried  up  above  ground. 

The  cotyledons  are  oblong-ovate,  sharply  auriculate,  short- 
petiolate.  The  blades  are  quite  thick.  When  they  first  emerge 
from  the  seed  coat  they  are  7-8  mm.  long  but  are  finally  10 
mm.  long  and  6  mm.  broad.  The  hypocotyl  is  rather  stout, 
10—30  mm.  long. 

The  foliage  leaves  are  ovate,  entire,  petiolate.  The  first  two 
are  opposite  or  nearly  so,  later  ones  are  alternate.  The  epi- 
cotyl is  short,  not  usually  more  than  2  or  3  mm.  in  length  when 
the  first  leaves  are  well  developed.  It  eventually  may  grow  to 
a  length  of  4-8  mm. 

The  seedlings  of  this  plant  resemble  very  much  those  of  E. 
angustifolia  microcarpa*  save  that  in  the  latter  the  petioles  of 
the  cotyledons  are  much  longer. 

MYRTACE^E. 

Eucalyptus  globulus  LABILL. 

This  is  the  well-known  "  blue  gum  "  tree  of  Australia.  It 
is  planted  extensively  in  California.  The  seeds  germinate  in 
from  one  to  two  weeks.  The  seed  coat  is  often  carried  up  by  the 
cotyledons.  These  are  doubled  over  each  other.  One  lobe  of 
each  is  exposed. 

The  hypocotyl  is  slender,  about  30  mm.  long.  The  cotyle- 
dons, when  fully  opened,  are  short-petiolate,  3  mm.  long  and 
generally  twice  as  broad,  two-lobed,  the  sinus  shallow.  When 
first  out  of  the  seed  coat  the  cotyledons  are  about  one-half  the 
size  here  named.  No  distinct  venation  was  observed,  although 
Lubbockf  states  that  they  are  tri-nerved. 

The  epicotyl  is  about  10  mm.  long.  The  foliage  leaves  are 
opposite,  lanceolate  and  entire,  those  higher  on  the  stem  be- 

*Lubbock,  op.  cit.  2  :  465. 
tOp.  cit.   I  :  530. 


Ramaley :     SEEDLINGS  OF  CERTAIN  WOODY  PLANTS.          81 

coming  gradually  broader.     Higher  internodes  of  the  stem  are 
quadrangular. 

Eucalyptus  citriodora  HOOK. 

The  mode  of  germination  and  the  seedling  of  the  "lemon- 
scented  gum"  resemble  the  species  just  described.  There  are 
some  important  points,  however,  to  be  noted. 

The  hypocotyl  is  20  mm.  long  and  quite  slender.  The  coty- 
ledons are  petiolate.  The  blade  is  broadly  orbicular,  entire,  in- 
distinctly 3-veined ;  at  length  6-9  mm.  broad,  4-7  mm.  long, 
green  above,  red  to  purple  below.  The  petiole  is  3-4  mm. 
in  length.  The  cotyledons  are  persistent  for  a  considerable 
time ;  often  remaining  till  ten  or  more  nodes  of  the  stem  are 
developed. 

Eucalyptus  corymbosa  SM. 

This  plant,  also  a  native  of  Australia,  is  called  "  blood-wood." 
Seeds  germinate  in  two  or  three  weeks.  The  hypocotyl  is  20 
mm.  long  and  quite  slender.  The  cotyledons  are  short-petiolate. 
The  blade  is  reniform,  deeply  cordate  at  base,  at  first  2-3 
mm.  long  and  5-6  mm.  broad.  It  finally  grows  about  twice 
this  size  and  is  indistinctly  3-veined. 

CORNACE.E. 

Cornus  amomum  MILL. 

The  seeds  of  the  common  "  dogwood  "  germinate  in  two  or 
three  weeks  after  planting,  but  sometimes  not  till  the  follow- 
ing year. 

The  hypocotyl  is  rather  slender  and  quite  long,  usually  50 
mm.  or  more  in  length.  The  epicotyl  also  is  greatly  elongated, 
reaching  a  length  of  40  mm.  The  cotyledons  are  oblong-ellip- 
tical, entire,  short-petiolate.  At  first  they  are  10  mm.  long  and 
5  mm.  broad.  The  blades  become  20  mm.  long  and  10  mm. 
broad,  the  petioles  4  mm.  long. 

Leaves  are  all  opposite,  ovate,  acute,  petiolate.  The  first  are 
like  the  later  ones. 

Cornus  stolonifera  MICHX. 

Seedlings  of  this  plant  resemble  those  of  the  previous  species 
in  all  essential  respects. 


82  MINNESOTA    BOTANICAL    STUDIES. 

Cornus  florida  LINN. 

Seedlings  of  the  "flowering  dogwood"  resemble  those  of  C. 
amomum. 

STYRACACE^:. 
Mohrodendron  carolinum  (LiNN.)  BRITT. 

Seeds  of  this  plant,  the  "snow-drop  tree,"  planted  in  the 
spring  of  the  year  following  their  ripening  lie  dormant  an  entire 
year  before  germinating. 

The  cotyledons  are  thin,  oval-oblong  in  outline,  rather  short- 
petiolate.  The  blades  are  at  first  20  mm.  long  and  8  mm. 
broad.  They  do  not  increase  much  in  size.  The  hypocotyl  is 
stout,  from  25-35  mm.  long. 

The  epicotyl  is  about  20  mm.  long.  The  leaves'are  all  alter- 
nate, ovate-acute,  serrate,  petiolate.  Save  in  size  there  is  no 
difference  between  the  first  and  the  later  leaves. 

BIGNONIACE^:. 
Tecoma  radicans  (LINN.)  DC. 

This  is  a  woody  climber,  the  "  trumpet  creeper,"  indigenous 
to  eastern  North  America  and  frequently  cultivated.  The  seeds 
germinate  in  about  ten  days.  The  large  flat  wing  of  the  seed 
is  sometimes,  though  not  usually,  carried  up. 

The  cotyledons  are  broadly  orbicular  and  deeply  notched  at 
the  apex.  They  are  almost  sessile.  When  first  above  ground 
they  are  5  mm.  wide,  but  when  fully  open  are  9  mm.  wide. 
They  do  not  increase  in  size  after  that  time.  The  hypocotyl  is 
20—30  mm.  long,  green  or  pale,  sometimes  pinkish. 

The  epicotyl  is  at  first  quite  short,  but  lengthens,  when  the 
foliage  leaves  open,  to  about  15  mm.  The  first  leaves  are 
simple,  ovate,  dentate,  petiolate,  distinctly  veined.  The  next 
leaves  are  tri-foliate.  Leaves  at  length  are  pinnately  com- 
pound. 

Catalpa  speciosa  WARDER. 

This  large  tree  is  a  native  of  the  southern  United  States. 
Seeds  germinate  in  from  one  to  two  weeks.  The  flat  winged 
seedcoat  is  sometimes  carried  up,  but  more  usually  remains  in 
the  soil. 

The  cotyledons  are  face  to  face.      They   are   dark  green, 


Ramaley:     SEEDLINGS  OF  CERTAIN  WOODY  PLANTS.          83 

deeply  bifid,  the  lobes  more  or  less  obovate,  5-6  mm.  long 
and  3-4  mm.  broad.  They  increase  rapidly  to  nearly  twice 
their  original  size.  The  hypocotyl  is  stout,  30  mm.  long. 

The  epicotyl  is  8-12  mm.  long.  Foliage  leaves  are  oppo- 
site, entire,  pointed,  ovate  to  cordate,  petiolate  with  distinct 
veining. 

Seedlings  of  this  plant  have  been  previously*  described,  but 
without  measurements  or  illustrations. 

RUBIACEJE. 

Cephalanthus  occidentalis  LINN. 

The  "button  bush"  is  a  low  shrub  indigenous  throughout 
most  of  North  America.  The  seed  germinates  in  about  three 
weeks.  The  seed  coat  remains  in  the  ground. 

The  cotyledons  are  ovate,  acute,  short-petiolate,  3  mm.  long 
and  i  mm.  broad  when  they  first  appear ;  at  length  they  become 
about  twice  or  three  times  that  size.  The  hypocotyl  is  slender, 
15-30  mm.  in  length. 

When  the  first  foliage  leaves  are  open  the  epicotyl  is  from 
4-8  mm.  long.  Leaves  are  opposite,  ovate,  acute,  entire,  long- 
petioled,  distinctly  veined. 

CAPRIFOLIACE^E. 

Sambucus  pubens  MICHX. 

This  is  the  "  red-berried  elder  "  of  the  northern  United  States. 
The  seeds  ripen  in  June.  If  sown  at  once  they  germinate  in 
about  one  month.  Some  of  the  seeds,  however,  do  not  come 
up  until  the  following  spring. 

The  hypocotyl,  which  passes  gradually  into  the  root,  is  about 
10  or  15  mm.  long.  The  cotyledons  are  petiolate.  When  they 
first  appear  they  are  3  mm.  long  and  2  mm.  broad.  The  coty- 
ledons become  longer  petioled  and  the  blades  more  ovate  as  they 
grow  older.  By  the  time  two  pairs  of  foliage  leaves  have  ap- 
peared they  are  10-15  mm-  in  length  with  petiole  8  mm.  long. 

The  epicotyl  is  very  short  as  are  also  the  succeeding  inter- 
nodes.  Leaves  are  opposite ;  the  first  two  pairs  cordate,  ser- 
rate, with  petioles  as  long  as  the  blades.  The  next  leaves  are 
generally  trifoliate ;  later  ones  are  pinnately  multifoliate. 

*Lubbock,  op.  cit.  2  : 335. 


84  MINNESOTA    BOTANICAL    STUDIES. 

GENERAL  OBSERVATIONS  ON  THE  FACTS    RECORDED  IN  THE 
PRECEDING  PAGES. 

Without  any  attempt  at  ecological  explanations  of  the  phe- 
nomena of  the  growth  and  development  of  seedlings  such  as 
given  by  Goebel*  a  few  generalizations  may  be  made  from  the 
plants  at  present  examined.  Some  of  the  features  to  which  at- 
tention is  called  have  been  previously  discussed  by  Klebsf  and 
LubbockJ  so  that  what  follows  will  not  be  so  much  a  considera- 
tion of  such  points  but  rather  a  classification  of  the  plants 
studied  with  regard  to  their  special  peculiarities. 

A  knowledge  of  the  shape  and  general  structure  of  the  coty- 
ledons does  not  help  one  to  predict  the  character  of  the  foliage 
leaves.  Sometimes  there  is  a  certain  resemblance  between  coty- 
ledons and  the  first  foliage  leaves  or  even  the  later  ones.  The 
resemblance  is,  however,  chiefly  in  cases  where  the  cotyledons 
are  ovate  or  oblong.  This  is  a  very  common  form  for  foliage 
leaves  as  well.  Thus  in  Toxylon  j>omiferum  and  Cephalanthus 
occidentals  the  cotyledons  and  foliage  leaves  are  much  alike, 
That  the  two  kinds  of  leaves  are  of  the  same  general  shape, 
may  be  a  mere  coincidence  and  of  no  great  significance. 

Where  the  general  shape  of  cotyledons  and  first  foliage  leaves 
is  much  the  same,  the  former  may  have  entire  margins  and  the 
latter  be  variously  toothed  or  lobed,  e.  g.,  Vitis  cordifolia,  Ptelea 
trifoliata.  While,  as  has  been  said,  there  is  no  absolute  agree- 
ment in  the  shape  of  cotyledons  in  a  given  genus  or  family, 
nevertheless,  there  are,  as  is  well  known,  many  families  in 
which  certain  types  of  cotyledons  prevail.  The  first  foliage 
leaves,  however,  are  more  frequently  alike,  e.  g".,  Acer  spp. 

In  cases  where  leaves  of  old  plants  are  pinnately  compound 
the  first  few  foliage  leaves  are  often  simple,  e.  g.,  Acer  negundo, 
Amor-pha  spp.,  Ptelea  trifoliata^  Schinus  molle,  Robinia  -pseuda- 
cacia,  Sambucus  spp.  In  all  these  cases  the  transition  to  the 
compound  form  is  gradual.  Thus  in  Ptelea  trifoliata  the  first 
leaf  is  simple,  the  second  leaf  usually  has  but  one  lateral  leaf- 
let. In  Robinia  -pseudacacia  the  second  leaf  is  trifoliate  while 
later  leaves  are  more  and  more  multifoliate. 

Occasionally  even  the  first  foliage  leaf   is  compound,  as  in 

*  Organographie  der  Pflanzen,  1898. 

tBeitrage  zur  Morph.  und  Biol.  der  Keimung.     Pfeffer's  Untersuchungen  aus 
dem  Botan.  Inst.  zu  Tubingen  i:  536.     1885. 
£Op.  cit. 


Ramaley :     SEEDLINGS  OF  CERTAIN  WOODY  PLANTS.          85 

Parthenocissus  guinquefolia.  In  Ailanthus  glandulosa^  however, 
the  first  few  leaves  are  merely  trifoliate  while  later  ones  are  pin- 
nate. Parkinsonia  and  Glcditsia  produce  pinnate  foliage  leaves 
at  once,  although  the  earlier  leaves  have  fewer  leaflets  than  those 
that  come  afterward. 

If  the  later-formed  leaves  are  not  compound  but  merely 
lobed  or  cleft  there  may  be  traced  a  more  or  less  gradual  transi- 
tion to  that  shape  from  the  entire  or  more  nearly  entire  first 
leaves,  e.  g.,  Brottssonetia  papyri/era,  Liriodendron  tulipifera. 

In  nearly  all  cases  where  the  first  two  leaves  are  opposite  and 
the  later  ones  alternate,  it  is  to  be  noted  that  the  third  and  fourth 
are  nearly  opposite,  the  fifth  and  sixth  are  closer  together  on 
the  stem  than  the  fourth  and  fifth  or  than  the  sixth  and  seventh  ; 
e,  g.,  Rhamnus  purshiana,  Eucalyptus  spp.,  Ulmus  spp.  In 
other  words,  the  transition  from  the  opposite  to  the  alternate  ar- 
rangement is  usually  gradual. 

The  cotyledons  of  many  species  increase  considerably  in  size 
after  they  escape  from  the  seed  coat ;  this  is  particularly  notice- 
able in  Schimu  molle,  Cercis  canadensis,  and  some  others.  In 
other  species  there  is  very  little  increase  in  the  size  of  the  coty- 
ledons after  they  first  appear,  e.  g.,  Rhamnus  purshiana,  Ailan- 
thus  glandulosa. 

Cotyledons  of  rather  remarkable  shape  were  noted  in  the  fol- 
lowing species :  Celtis  occidentalism  Catalpa  spcciosa,  Euca- 
lyptus globulus,  Tecoma  radicans,  Acer  negundo,  Berchemia 
racemosa,  Butncria  florida  and  fertilis.  The  first  four  named 
have  the  cotyledons  bifid  or  variously  notched  or  retuse. 

Catalpa  and  Tecoma,  both  Bignoniaceous  plants,  have  very 
similar  cotyledons.  The  peculiar  asymmetrical  cotyledons  of 
Butneria  florida  are  reproduced  exactly  in  B.  fertilis.  Euca- 
lyptus globulus,  on  the  other  hand,  does  not  agree  at  all,  in  the 
shape  of  its  cotyledons,  with  E.  citriodora  and  E.  corymbosa. 
These  have  rotund-orbicular  cotyledons.  The  long,  narrow 
cotyledons  of  Acer  negundo  are  quite  different  from  those  of 
A.  saccharinum.  Berchemia  racemosa  has  ligulate  cotyledons, 
while  in  Rhamnus  -purshiana,  the  only  other  plant  of  the  same 
family  investigated,  the  cotyledons  are  obovate.  The  large 
notched  cotyledons  of  Celtis  occidentalis  do  not  resemble  those 
of  the  other  Ulmaceae  examined.  This,  is,  however,  to  be  ex- 
pecied  from  the  great  difference  in  the  character  of  the  fruit 
in  Celtis  and  Ulmus. 


86  MINNESOTA    BOTANICAL    STUDIES. 

From  the  foregoing  it  may  be  concluded  that  broad  general- 
izations in  regard  to  the  shape  of  cotyledons  in  plant  families, 
cannot  be  safely  made  without  a  considerable  mass  of  data. 

EXPLANATION  OF  PLATES. 

Plate  I.  Seedlings  in  various  stages  of  the  following  plants :  Popu- 
lus  deltoides,  Ulmus  americana,  Celtis  occidentalis,  Toxylon  pomi- 
ferum,  Broussonetia  papyrifera,  Liriodendron  tulipifera,  But- 
neria  florida,  Parkinsonia  aculeata. 

Plate  II.  Seedlings  in  various  stages  of  the  following  plants  :  Cercis 
canadensis,  Amorpha  fruticosa,  Amorpha  nana,  Robinia  psetidaca- 
cia,  Ptelea  trifoliata,  Ailanthus  glandulosa,  Schinus  molle,  Celas- 
trus  scandens. 

Plate  III.  Seedlings  in  various  stages  of  the  following  plants:  Acer 
negundo,  Acer  saccharimim,  Acer  glabrum,  Berchemia  racemosa, 
Rhamnus  purshiana,  Vitis  cordifolia,  Parthenocissus  quinquefolia, 
Sterculia  plantanifolia. 

Plate  IV.  Seedlings  in  various  stages  of  the  following  plants : 
Elceagnus  ttmbellata,  Eucalyptus  globulus,  Eucalypttis  citriodora, 
Cornus  amomum,  Afohrodendron  carolinum,  Catalpa  speciosa,  Te- 
coma  radicans,  Cephalanthus  occidentalis,  Sambucus  pubens. 

The  amount  of  enlargement  or  reduction  is  indicated  for  each  plant. 


VOL.   II. 


MINNESOTA 


Populus  deltoides  x 


Ulmus  amerie.ana  x 


Broussonetia  papyrifera  x 


Liriodendroh  tulipifera  (natural 


[CAL    STUDIES. 


PART    II. 


Coltis  occidentnlis  x  .•{ 


Toxylon  pomiferum  x  {( 


Butneria  florida  x 


Pnrkinsonin  aculeata  x 


VOL.    II. 


MINNESOT 


Amorpha  fruticosa  x 


Ptelea  trifoliata  x 


Ailanthus  glai 


STUDIES. 


PART    II 


Amorpha  nana  x 


Robinia  pseudacacia  x  f 


\ 


Schinus  molle  x  | 


Celastrus  scanuons  x 


VOL.    II. 


MINNES 


Acer  negundo  x  £ 


Acer  saccharinum  x  f 


\ 


o 


l\ 

Rhamnus  purshiana  x 


Vitis  cordifolia  x  J 


:AL  STUDIES. 


PART    I 


Acer  glabrum  x  £ 


Parthenocissue  quinquefolia  x 


Sterculia  platanifolia  x  \ 


[I. 


VOL.    II. 


MINNESOTA    BC 


Elaeagnus  umbellata  x 


Eucalyptus  corymbosa 
(natural  size) 


Eucalyptus  globulus 
(natural  size) 


Eucalyptus  citriod 
(natural  size) 


Catalpa  speciosa  x  £ 


Tecoma  radicans 


CAL    STUDIES. 


PART    II 


K 


Cornus  amomum  x  £ 


Mohrodendron  carolinum  x  $ 


Cephalanthus  occidental  x  1$  Sambucus  pubens  x  J 


V. 


IX.  COMPARATIVE  ANATOMY  OF  HYPOCOTYL 
AND  EPICOTYL  IN  WOODY  PLANTS. 

FRANCIS  RAMALEY. 

The  following  is  an  account  of  the  anatomy  of  seedlings  of 
certain  woody  dicotyledonous  plants.  These  plants  were  studied  : 
Ulmus  amcricana  LINN.,  Celtis  occidentalis  LINN.,  Toxylon 
pom  if cr  um  RAF.,  Broussonctia  papyrifera  (LiNN.)  VENT.,  Li- 
riodcndron  tiilipifcra  LINN.,  Menispermum  canadense  LINN., 
Butneria jlorida  (LINN.)KEARNEY,  Parkinsonia  aculeataLiNTx., 
Cercis  canadensis  LINN.,  Gleditsia  triacanthos  LINN.,  Amorpha 
fruticosa  LINN.,  Robinia  pseudacacia  LINN.,  Ptelea  trifoliata 
LINN.,  Ailanthus  glandulosa  DESF.,  Schinns  molle  LINN.,  Ber- 
chemia  racemosa  SIEB.  &  Zucc.,  Rhamnus  purshiana  DC., 
Vitis  cordifolia  NLicuyL^EZaeagmts  umbellata  THUNB.,  Eucalyp- 
tus globulus  LABILL.,  Tecoma  radicans  (LiNN.)  DC.,  Ca- 
talpa  speciosa  WARDER,  Cephalanthus  occidentalis  LINN.  The 
order  in  which  they  are  described  is  that  of  Engler  and  Prantl. 
This  order  will  be  followed  throughout. 

The  author  is  under  obligation  to  Professor  Conway  MacMil- 
lan,  who  suggested  the  subject  of  the  investigation  and  under 
whose  direction  the  work  has  been  completed. 

The  seedlings  were  grown  at  the  University  of  Minnesota 
during  the  years  1896,  1897  and  1898.  They  were  examined  at 
different  ages  so  that  the  original  structure  of  both  hypocotyl 
and  epicotyl  could  be  noted  as  well  as  the  differences  brought 
about  through  secondary  changes. 

For  the  sake  of  convenience  and  uniformity  three  stages  were 
studied ;  these  may  be  designated  as  first,  second  and  third 
stages.  A  seedling  with  the  cotyledons  expanded  but  with  the 
epicotyl  undeveloped  is  said  to  be  in  the  first  stage.  Obviously 
only  the  structure  of  the  hypocotyl  was  studied  in  this  stage. 
In  the  second  stage  the  epicotyl  has  elongated  and  the  first  foli- 
age leaves  have  appeared.  In  the  third  stage  a  considerable 
number  of  foliage  leaves  have  been  developed  and  the  anatom- 
ical structure  has,  to  a  considerable  extent,  taken  on  its  perma- 


88  MINNESOTA    BOTANICAL    STUDIES. 

nent  characters.  Sections  were  also,  in  many  cases,  cut  from 
material  two  years  old  for  purposes  of  comparison. 

Since  the  structure  of  the  hypocotyl  is  often  materially  differ- 
ent in  all  the  three  mentioned  stages,  it  has  seemed  important  to 
make  a  record  of  the  changes  which  take  place  during  the  first 
year's  growth.  Previous  investigators  have  not  done  this. 

A  number  of  investigators  who  have  made  a  comparative  study 
of  root  and  shoot  have  incidentally  examined  the  hypocotyl, 
e.g.,  Goldsmith  [1876]  and  Gerard  [1880  and  1881].  The 
latter  made  some  careful  observations  on  the  course  of  vascular 
bundles  from  the  cotyledons  to  the  root.  His  statement  that  the 
characteristic  root  structure  often  extends  as  high  as  the  coty- 
ledons is  not,  in  general,  confirmed  by  the  present  investigation. 

The  most  important  articles*  which  need  to  be  mentioned  at 
the  present  time  are  by  Dangeard  [1888  and  1889],  Van  Tieg- 
hem  [1891],  and  Flot  [1889  and  1890].  Dangeard  begins  with 
a  study  of  the  structure  of  roots,  of  which  he  distinguishes  three 
types.  In  the  first  type  the  root  is  diarch ;  the  hypocotyl  has 
four  bundles  in  two  pairs  which  arise  as  cotyledonary  trace  bun- 
dles by  the  division  of  the  midrib  of  each  cotyledon.  In  the 
second  type  the  root  is  tetrarch ;  the  hypocotyl  has  eight  bun- 
dles in  four  groups.  In  the  third  type  the  root  is  octarch,  while 
the  hypocotyl  has  sixteen  bundles  in  eight  groups.  The  first 
type  of  structure  of  the  hypocotyl  above  mentioned  is  the  one 
commonly  found  in  the  plants  studied  by  the  present  writer  who 
has  called  it  the  "  typical  structure."  (See  General  Conclu- 
sions at  the  close  of  this  paper.) 

Flot  [1889,  1890]  describes  the  "region  tigellaire,"  a  much 
thickened  portion  of  the  axis  of  certain  year-old  seedlings.  The 
region  extends  from  the  base  of  the  hypocotyl  up  to  the  first 
foliage  leaf  or  to  some  point  between  that  and  the  cotyledons. 
It  is  noted  only  in  certain  species.  It  is  not  the  same  as  the 
"  tigelle,"  which  extends  only  as  high  as  the  cotyledons.  The 
"region  tigellaire"  is  characterized  by  only  a  slight  develop- 
ment of  sclerenchyma  and  of  normal  phloem,  while  internal 
phloem  is  probably  altogether  absent.  The  pericycle,  he  says, 
is  well  developed. 

Van  Tieghem  [1891]  divides  the  hypocotyl  into  "tigelle" 
and  "  rhizelle."  The  growth  of  the  hypocotyl  is  produced  by 
the  elongation  of  either  the  tigelle,  as  in  Ricinns,  Acer,  Cucur- 

*  Search  has  been  made,  but  without  success,  for  a  paper  by  Monal :  Rech. 
sur  1'anat.  compar.  de  la  tige  hypocot.  et  epicot. 


Ramaley  :   HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.       89 

bita,  Tagetes,  Convolvulus  and  Mirabilis,  or,  the  rhizelle,  as  in 
Ranunculaceae,  Cruciferae,  Caryophyllaceas,  Chenopodiaceae, 
Umbelliferse,  Rubiaceaa  and  Coniferas,  or  by  a  combination  of 
the  growth  of  both  as  in  Euonymus. 

The  designation  of  certain  regions  as  tigelle,  rhizelle  and 
tigellaire  does  not  seem  to  the  present  writer  a  matter  of  great 
importance  in  the  plants  which  he  has  studied,  for  in  them  these 
regions  are  by  no  means  sharply  differentiated.  Further  obser- 
vations and  references  to  the  work  of  Plot  mentioned  above 
are  given  in  the  pages  which  follow. 

In  the  special  portion  of  the  present  work  will  be  found  de- 
scriptions of  the  structure  of  hypocotyl  and  epicotyl  in  the 
various  species  examined.  Accompanying  each  description  is  a 
diagram  of  the  cross  section  of  the  hypocotyl  when  the  seedling 
is  in  the  first  stage  previously  described,  and  diagrams  of  both 
hypocotyl  and  epicotyl  of  the  second  and  third  stages.  In  these 
diagrams  stereom  is  black,  xylem  is  dotted,  cortex,  phloem, 
pericycle  and  the  pith  are  white.  The  endodermis,  when  dis- 
tinct, is  indicated  by  a  single  line  as  is  also  the  epidermis  and 
the  boundaries  between  the  various  zones.  In  each  figure  the 
diagrams  of  the  hypocotyl  are  at  the  left,  those  of  the  epicotyl 
at  the  right. 


Ulmus  americana. 
Structure  of  Hypocotyl. 

The  epidermis  is  composed  of  cells  which,  in  cross  section, 
are  square  or  rounded.  After  secondary  growth  of  the  stele  has 
commenced  these  cells  become  very  much  flattened.  There  is 
no  hypoderma  differentiated.  The  cells  of  the  cortex  are  large  ; 
all  are  about  the  same  size. 

The  endodermis  is  small-celled  and  is  easily  recognized  in 
early  stages,  when  it  contains  very  little  starch.  Afterward 
starch  becomes  abundant  in  the  endodermis,  pericycle,  cortex, 
phloem  and  inner  xylem. 

In  the  stele  there  are  many  small  phloem  bundles  which  are 
confluent  into  two  crescent-shaped  areas.  There  are  two  xylem 
bundles  of  somewhat  crescentic  appearance  in  cross  section. 
The  xylem  and  phloem  soon  form  closed  rings. 

The  pericycle,  in  seedlings  which  have  about  two  internodes 


90 


MINNESOTA    BOTANICAL    STUDIES. 


of  the  stem  developed,  is  partially  sclerenchymatous.  At  a  later 
time  numerous  groups  of  sclerenchyma  are  found  in  the  phloem 
and  cortex. 

The  pith  becomes  quite  small.  There  is  a  small-celled  peri- 
medullary  zone. 

The  formation  of  cork  cambium,  as  noted  by  Plot  (  [1890],  p. 
29  ),  takes  place  in  the  inner  cortex. 

Structure  of  EpicotyL 

The  cells  of  the  epidermis  when  seen  in  cross  section,  are 
somewhat  rectangular  in  outline.  The  tangential  diameter  is 
the  longer.  Numerous  hairs  are  present.  No  collenchymatous 
hypoderma  is  produced.  The  cortex  is  rather  narrow.  The 
cells  are  all  about  the  same  size. 

The  endodermis  is  distinct  only  in  young  material.  The  cells 
are  small.  They  contain  starch.  At  the  end  of  the  season 
starch  is  found  in  the  pith  and  inner  xylem  and  is  sparingly  dis- 
tributed in  the  cortex  and  phloem. 

In  the  youngest  material  examined  the  phloem  forms  a  closed 
ring  surrounding  a  number  of  xylem  groups.  There  are  usually 
eight  of  these.  They  soon  fuse  to  form  a  complete  ring. 

Numerous  small  groups  of  thick-walled  cells  finally  make 
their  appearance  in  cortex,  pericycle  and  phloem. 

The  pith  becomes  quite  small.     The  cells  have  thin  unligni- 

fied  walls.  The  perimedullary  zone 
is  easily  distinguished ;  it  consists  of 
from  one  to  three  layers  of  small  cells 
which  are  often  somewhat  flattened. 

Cork  formation,  as  is  well  known 
in  this  species,  begins  in  the  outer- 
most cell  layer  of  the  cortex. 

Comparison  of  Structure  of  Hypo- 

cotyl  and  EpicotyL 
The  epidermal  cells  of  the  hypo- 
cotyl  in  young  material  appear  radi- 
ally elongated,  those  of  the   epicotyl 
tangentially  elongated.     The  former 
region  has  a  thicker  cortex,  fewer  epi- 
dermal hairs,  sclerenchyma  developed 
FIG.  i.  earlier  in  the  pericycle. 


Ulmus 

americana 


Ramaley :    HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.       91 

The  stele  of  the  hypocotyl  has  two  xylem  bundles  and  two 
aggregations  of  phloem  bundles.  In  the  epicotyl  the  youngest 
material  examined  has  a  complete  ring  of  phloem  and  about 
eight  xylem  bundles.  Cork  formation  in  the  former  region 
takes  place  deep  in  the  cortex  instead  of  in  the  outermost  corti- 
cal layer. 

In  their  final  structure  the  two  regions  are  practically  alike. 

Celtis  occidentalis. 

Structure  of  Hypocotyl. 

The  epidermis  is  composed  of  thin-walled  cells,  small,  square 
in  cross  section.  There  is  no  hypoderma.  The  elements  of 
the  cortex  are  large.  There  are  about  twenty  layers  of  cells. 

The  cells  of  the  endodermis  are  much  smaller  than  those  of 
the  cortex  and  on  this  account  the  endodermis  is  readily  distin- 
guished until  considerable  secondary  growth  of  vascular  tis- 
sue has  taken  place. 

Starch  is  found  in  the  endodermal  region  from  the  first ; 
toward  the  close  of  the  season  it  is  found  not  only  in  the  pith, 
phloem  and  cortex,  but  very  abundantly  distributed  throughout 
the  xylem.  Large  isodiametric  crystals,  long  known  in  the 
stem  of  this  species  (Moeller  [1882],  p.  74),  make  their  appear- 
ance in  the  cortex  some  time  before  the  close  of  the  first  season. 

The  stele,  which  is  cylindrical  from  the  first,  has  originally 
four  xylem  bundles  and  two  crescentic  masses  of  phloem.  At 
an  early  stage  the  xylem  forms  a  closed  ring,  while  it  is  not 
till  sometime  afterward  that  the  two  areas  of  the  phloem  become 
united. 

Two  interrupted  rings  of  sclerenchyma  appear  later  in  the 
first  season,  one  of  these  is  in  the  cortex  and  consists  of  much 
larger  groups  of  cells  than  does  the  other  which  is  in  the  outer 
phloem. 

The  pith  is  large-celled.  A  more  or  less  definite  perimedul- 
lary  zone  of  small  cells  is  at  length  developed. 

Cork  formation  begins  at  a  late  period  in  the  outermost  cell 
layer  of  the  cortex. 

Structure  of  Epicotyl. 

The  cells  of  the  epidermis  are  at  first  nearly  square  in  cross 
section  but  at  a  later  time  are  considerably  flattened.  There 


92 


MINNESOTA    BOTANICAL    STUDIES. 


are  numerous  simple  curved  and  pointed  hairs ;  there  are  also 
some  with  bulbous  ends. 

The  cortex  is  thin.  A  distinct  collenchymatous  hypoderma 
is  developed.  It  usually  consists  of  three  or  four  layers  of 
cells. 

The  endodermis,  which  is  originally  distinct,  soon  becomes 
unrecognizable.  The  cells  are  about  the  same  size  as  those  of 
the  cortex ;  they  contain  starch.  Eventually  all  the  parenchy- 
matous  elements  contain  starch. 

The  stele  is  originally  somewhat  elliptical  in  cross  section. 
The  phloem,  in  the  youngest  material  examined,  forms  a  com- 
plete ring.  There  are  generally  two  large  and  four  small 
xylem  bundles.  These  soon  fuse  to  form  a  closed  xylem 
zone. 

An  interrupted  band  of  sclerenchyma  is  developed  at  the 
outer  limit  of  the  xylem. 

As  in  Celtis  australis  (cf.  Flot  [1893],  p.  68)  there  is  a  dis- 
tinct perimedullary  zone  composed  of  two  or  three  cell  rows. 

Cork  formation  begins,  rather  late  in  the  season,  in  the  outer- 
most hypodermal  layer  (cf.  Moeller  [1882],  p.  74). 


Comparison  of  Structure  of  Hypocotyl  and  E-picotyL 

A  striking  difference  between  hypocotyl  and  epicotyl  is  the  ab- 
sence from  the  former  region  of  the 
numerous  epidermal  hairs  so  abun- 
dant in  the  latter.  The  hypocotyl 
is  without  a  hypoderma. 

The  primary  stelar  structure  of 
the  hypocotyl  is  peculiar,  the  phloem 
forming  two  crescentric  masses  and 
not  uniting  into  a  closed  ring  till 
after  the  xylem  bundles  have  fused. 
The  epicotyl  possesses  a  ring  of 
phloem  and  six  xylem  bundles. 

At  the  end  of  the  season  the  hypo- 
cotyl has  two  interrupted  bands  of 
sclerenchyma  instead  of  one,  and  a 
smaller  pith.  Aside  from  these  dif- 
ferences the  two  regions  are  the 
FIG.  2.  same  in  structure. 


Celtis 

occidentalis 


Ramaley :   HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.       93 

HORACES. 

Toxylon  pomiferum. 

Structure  of  Ifypocotyl. 

In  cross  section  the  cells  of  the  epidermis  appear  radially 
elongated.  Eventually  they  are  considerably  flattened.  The 
cells  of  the  outer  cortex  are  similar  to  those  of  the  epidermis. 
There  is  no  collenchyma  developed.  The  inner  cortex  is  com- 
posed of  larger  cells. 

The  endodermis  is  distinct  but  in  material  taken  at  the  close 
of  the  growing  season  it  was  not  distinguished.  The  develop- 
ment of  pericycle  is  remarkable.  This  region  is  composed  of 
about  six  layers  of  parenchymatous  cells  resembling,  in  shape, 
those  of  the  endodermis. 

Starch  is  found,  from  the  first,  in  the  endodermis  and  later 
appears  in  all  the  conjunctive  tissues. 

The  stele  is  slightly  four-angled.  There  are,  in  the  young- 
est material  examined,  four  xylem  bundles  and  two  large  cres- 
cent-shaped phloem  bundles.  The  phloem  soon  forms  a  com- 
plete ring  as  does  also  the  xylem,  but  the  two  xylem  bundles 
first  fuse  in  pairs.  - 

About  this  time  four  aggregations  of  small  groups  of  scleren- 
chyma  appear  in  the  pericycle.  Eventually  a  nearly  complete 
sclerenchymatous  ring  surrounds  the  phloem. 

The  pith  is  large-celled.  A  small-celled  perimedullary  zone 
of  three  or  four  layers  is  present. 

Cork  formation  takes  place  in  the  fourth  or  fifth  layer  of  the 
cortex. 

Structure  of  EpicotyL 

The  epidermis  is  composed  of  cells  which  are,  at  first,  nearly 
square  in  cross  section  but  later  are  very  much  flattened.  Ac- 
cording to  Moeller  [1882]  the  epidermis  is  two-layered. 
Numerous  straight  epidermal  hairs  are  present ;  there  are  also 
some  stalked  glandular  hairs.  The  cells  of  the  cortex  are 
rather  small,  parenchymatous,  not  at  all  collenchymatous. 

The  endodermis  is  distinguished  with  difficulty  even  in  very 
young  material.  Its  cells  contain  starch.  Starch  is  later  found 
in  all  the  parenchymatous  tissues. 

There  is  a  variable  number  of  vascular  bundles ;  usually 
eight  to  sixteen.  These  soon  fuse  to  form  closed  rings  of 
phloem  and  xylem. 


94 


MINNESOTA    BOTANICAL    STUDIES. 


An  interrupted  sclerenchymatous  ring  is  formed  at  the  outer 
edge  of  the  phloem ;  the  cells  are  thick-walled  but  do  not  be- 
come lignified  the  first  year. 

The  pith  is  rather  large.  There  is  a  small-celled  perime- 
dullary  zone  which  is  quite  definite. 

Cork  formation  in  the  epicotyl  takes  place  in  the  outermost 
cortical  layer  (cf.  Moeller  [1882]  ). 

Comparison  of  Structure  of  Hypocotyl  and  Epicotyl. 

In  the  hypocotyl  the  cortex  and  pericycle  are  much  better 

developed  than  in  the  epicotyl.  The 
former  region  is  without  epidermal 
hairs.  This  point  of  difference  was 
previously  noted  by  Klebs  [1885]. 

The  stele  of  the  hypocotyl  has  orig- 
inally four  vascular  bundles,  instead  of 
from  eight  to  sixteen ;  the  pith  is  small 
in  extent. 

The  sclerenchyma  is  first  formed  in 
four  patches  but  afterwards  forms  al- 
most a  complete  ring.  Cork  formation 
in  the  hypocotyl  begins  in  a  deeper 
layer  of  the  cortex. 

At   the    close  of   the  year   the  two 
Toxyion  regions  have  nearly  the  same  structure, 

pomiferum   about  the  only  difference  being  the  size 
FIG.  3.  of  the  pith. 

Broussonetia  papyrifera. 
Structure  of  Hypocotyl. 

There  is  an  epidermis  of  small  cells  nearly  square  in  outline 
when  seen  in  cross  section.  These  cells  become  greatly  elon- 
gated tangentially  as  the  tissues  within  increase  in  thickness. 
Short,  blunt,  unicellular  epidermal  hairs  are  numerous. 

The  cortex  is  composed  of  about  six  layers  of  large,  thin- 
walled  parenchymatous  elements  which,  like  the  epidermal 
cells,  become  stretched  toward  the  close  of  the  season. 

The  endodermis  is  small-celled.  It  sometimes  remains  dis- 
tinct till  nearly  the  close  of  the  first  season.  Starch  is  present 
in  the  endodermis,  but  absent  from  all  other  tissues  for  a  long 


Ramaley :    HYPOCOTYL  AND  EPICOTYI,  IN  WOODY  PLANTS.       95 

time.  It  eventually  appears  in  the  pericycle,  phloem,  medullary 
rays  and  inner  elements  of  the  xylem. 

The  stele  is  originally  very  small.  In  the  disposition  of  the 
vascular  tissues  this  plant  differs  from  all  others  examined  by 
the  writer.  In  cross  section  the  center  of  this  stele  is  seen  to 
be  occupied  by  an  elongated  area  of  xylem.  On  each  side  of 
this,  separated  by  a  small  amount  of  conjunctive  tissue,  is  a 
crescent-shaped  mass  of  phloem.  The  xylem  soon  forms  a 
somewhat  four-sided  mass,  and  is  surrounded  by  a  ring  of 
phloem.  The  xylem  at  length  becomes  circular,  and  the  sur- 
rounding phloem  increases  greatly  in  amount. 

There  is  but  slight  development  of  stereom,  although,  toward 
the  close  of  the  first  season,  numerous  isolated  sclerenchymatous 
elements  are  found  in  the  phloem. 

The  cork  cambium  originates  in  the  endodermis  or  pericycle. 
The  ring  of  phellogen  is  sometimes  irregular,  appearing  now 
in  one,  now  in  the  other  of  the  regions  named. 

It  may  be  said  that,  since  the  structure  of  the  hypocotyl  in 
this  species  so  much  resembles  the  general  type  of  root  struc- 
ture, it  was  thought  best  to  examine  a  large  number  of  plants, 
lest  the  peculiarities  noted  should  have  been  due  to  teratological 
development.  All  the  plants  were,  however,  found  to  be  alike. 
Neither  is  there  any  trouble  in  this  species,  to  determine  the 
lower  limit  of  the  hypocolyl,  for  it  is  enlarged  below  and  does 
not  gradually  shade  off  into  root,  as  is  the  case  in  some  seed- 
lings. 

Structure  of  Epicotyl. 

The  epidermis  is  small-celled.  There  are  numerous  simple, 
blunt  and  pointed  hairs,  and  also  some  with  a  single  stalk  cell 
and  a  multicellular  bulb  at  the  distal  end. 

A  somewhat  collenchymatous  hypoderma  is  developed,  con- 
sisting of  two  or  three  layers  of  cells,  which  are  smaller  than 
the  deeper  cells  of  the  cortex. 

The  small-celled  endodermis,  at  first  distinct,  soon  becomes 
displaced  and  changed,  owing  to  secondary  growth  of  sub-lying 
tissues. 

Starch  is  almost  entirely  absent,  except  in  the  endodermal 
region,  till  about  the  close  of  the  first  growing  season,  when  it 
appears  in  the  pith,  medullary  rays,  phloem  and,  to  a  slight 
extent,  in  the  cortex. 


96 


MINNESOTA    BOTANICAL    STUDIES. 


The  stele  is  large.  There  is  a  circle  of  twelve  to  eighteen 
conjoint  vascular  bundles.  These  soon  fuse  to  form  a  narrow 
zone  each  of  xylem  and  phloem. 

There  is  a  considerable  amount  of  sclerenchyma  at  the  outer 
edge  of  the  phloem.  The  cells  are,  however,  mostly  isolated  or 
else  occur  in  small  groups. 

The  pith,  which  is  extensive,  is  composed  of  large,  parenchy- 
matous  elements  with  thin,  slightly  lignified  walls.  According 
to  Flot  [1893],  there  is  a  perimedullary  zone  of  rive  or  six  lay- 
ers of  crushed,  thin-walled  cells.  The  same  author  states  that 
laticiferous  tubes  are  found  in  the  perimedullary  region  of 
young  twigs  of  this  species. 

The  cork  cambium  is  formed  in  the  outermost  hypodermal 
layer  (cf.  Moeller  [1882],  p.  82). 

Comparison  of  Structure  of  Hypocotyl  and  Epicotyl. 

Both  hypocotyl  and  epicotyl  have  simple  epidermal  hairs,  but 
the  former  does  not  have  the  pointed  or  the  bulbous  hairs  found 
in  the  latter  region.  The  hypocotyl  is  also  without  the  some- 
what collenchymatous  hypoderma 
found  in  the  epicotyl ;  its  endodermis 
persists  for  a  greater  length  of  time. 
The  structure  of  the  stele  in  the 
hypocotyl  is  anomalous.  A  single 
flat  bundle  of  xylem  is  flanked  by 
phloem,  which  eventually  surrounds 
the  centrally-lying  xylem,  the  inner 
cells  of  which  contain  starch.  There 
is  no  pith.  The  epicotyl,  on  the  other 
hand,  has  a  large  pith,  and  the  vas- 
cular bundles  are  originally  numer- 
ous. Starch  is  absent  from  the  xylem. 
Cork  formation  is  endodermal  or 
pericyclic  in  the  hypocotyl,  but  hypo- 
dermal  in  the  epicotyl. 


Broussonetia 
papyrifera 


FIG.  4. 

MAGNOLIACE.3L 
Liriodendron  tulipifera. 
Structure  of  Hypocotyl. 

The  epidermis   consists  of  cells  which   are   nearly  square  in 
cross  section  ;  at  first  they  are  very  much  bulged.     They  never 


Ranialey :   IIYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.       97 

become  flat.  The  two  or  three  layers  of  the  cortex  just  below 
the  epidermis  are  small-celled.  The  deeper  layers  are  very 
large-celled. 

The  endodermis  is  small-celled  and  easily  distinguished  in 
young  material,  but  is  eventually  displaced  and  is  not  distin- 
guishable. In  the  young  stages  starch  is  entirely  absent  from 
the  hypocotyl,  but  later  is  found  sparingly  distributed  through 
the  various  parenchymatous  tissues. 

The  stele,  which  is  originally  quadrangular,  has  four  vascu- 
lar bundles  arranged  in  pairs.  By  their  continued  growth 
zones  of  xylem  and  phloem  are  produced. 

About  the  time  that  a  complete  ring  of  xylem  has  been  formed 
four  masses  of  sclerenchyma  appear  in  the  pericyle.  Even- 
tually other  groups  of  pericyclic  cells  also  become  sclerotic. 
The  phloem  immediately  under  these  groups  is  better  developed 
than  at  other  places. 

The  pith  is  slightly  quadrangular.  The  cells  are  thin-walled. 
A  definite  perimedullary  zone  was  not  distinguished. 

The  cork  cambium  is  produced  in  the  outermost  layer  of  cor- 
tical cells. 

Structure  of  EfocotyL 

The  cells  of  the  epidermis,  when  seen  in  cross  section,  ap- 
pear square  or  tangentially  elongated.  A  narrow  collenchyma- 
tous  hypoderma  is  developed.  The  remaining  cells  of  the  cor- 
tex are  all  about  the  same  size. 

The  endodermis  is  distinct  in  young  material,  owing  to  the 
presence  of  starch  in  its  cells.  At  a  later  time  starch  is  distrib- 
uted in  small  amount  in  the  various  parenchymatous  tissues. 

The  number  of  primary  xylem  groups  in  the  stele  is  about 
six  or  eight.  Groups  of  phloem  are  somewhat  more  numerous. 
Closed  zones  of  xylem  and  phloem  are  produced  very  early. 

The  outer  phloem  has  many  groups  of  sclerenchymatous 
fibers.  These  groups  are  close  together,  separated  only  by 
medullary  rays.  A  small  amount  of  sclerenchyma  is  produced 
in  the  cortex. 

The  pith  is  rather  large,  and  composed  of  cells  with  thin,  un- 
lignified  walls.  No  perimedullary  zone  was  distinguished. 

Cork  is  developed  in  the  outermost  cell  layer  of  the  cortex 
(cf.  Moeller  [1882],  p.  229). 


98 


MINNESOTA    BOTANICAL    STUDIES. 


Liriodendron 
tulipifera 

FIG.   5. 


Comparison    of   Structure   of  Hypocotyl  and   Epicotyl* 

The  cortex  of  the  hypocotyl  is  much  thicker  than  that  of  the 
epicotyl.     The  former  region  has   no  hypoderma ;  it  has  four 

vascular  bundles  instead  of  six  or  eight 
or  more ;  the  sclerenchyma  first  ap- 
pears in  only  four  groups  and  at  no 
time  is  as  well  developed  as  in  the 
epicotyl. 

The  pith  of  the  hypocotyl  is  smaller 
than  that  of  the  epicotyl ;  it  is  some- 
what quadrangular  in  shape. 

MENISPERMACE^E. 


Menispermum  canadense. 

Structure  of  Hypocotyl. 

The  epidermis  consists  of  cells  which 
are  square  or  rectangular  in  cross  sec- 
tion. Late  in  the  first  season  they  be- 
come flat  and  tangentially  elongated. 
A  very  thick,  tough  cuticle  develops  at  the  same  time. 

There  is  no  hypoderma.  There  are  about  twelve  layers  in 
the  cortex.  The  cells  are  large. 

The  endodermis  consists  of  cells  smaller  than  those  of  the 
cortex.  It  remains  distinct  a  long  time,  but  was  not  distin- 
guished in  material  taken  at  the  close  of  the  growing  season. 
The  pericycle  is  peculiar.  It  is  one  or  two  layers  in  thickness. 
Usually  every  second  or  third  cell,  when  seen  in  cross  section, 
is  without  starch,  although  starch  is  present  in  the  other  cells. 
Eventually  these  cells  also  contain  starch. 

This  plant  is  somewhat  unique  in  the  distribution  of  starch  in 
its  tissues,  for  in  all  the  different  stages  examined  starch  was 
found  in  cortex,  pith,  endodermis  and  medullary  rays  and  in 
the  pericycle  except  as  just  noted. 

The  stele  is  quadrangular  and  has  four  primary  vascular 
bundles.  These  increase  considerably  in  size  as  the  plant 
grows  older.  At  the  close  of  the  growing  season  they  are  of 
about  the  same  extent  as  the  medullary  rays  which  are  com- 
posed of  wood  parenchyma  and  are  full  of  starch.  There  is  no 
phloem  produced  the  first  year  opposite  the  medullary  rays. 

The  pith  is  composed  of  large  cells.     There  is  a  rather  dis- 


Ramaley :   HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS. 


99 


tinct  smaller-celled  perimedullary  zone.     No  cork  is  formed  the 
first  year. 

Structure  of  Epicotyl* 

The  cells  of  the  epidermis,  at  first  square  in  cross  section,  be- 
come very  much  flattened  and  develop  a  thick  cuticle  like  that 
of  the  hypocotyl.  A  more  or  less  definite  collenchymatous  hy- 
poderma  is  developed.  The  cortex  is  composed  of  about  six 
cell  layers. 

The  endodermis  is  not  easily  distinguished  even  in  youngest 
stages.  The  pericycle  has  some  cells  which  in  cross  section 
appear  empty,  while  the  neighboring  cells  contain  starch. 
These  empty  cells  at  a  later  time  either  become  filled  with 
starch  or  else  are  displaced  so  that  they  are  not  recognized. 

Starch  is  present  in  the  cortex,  medullary  rays,  endodermis, 
pericycle  and  pith. 

There  are  originally  from  nine  to  fifteen  vascular  bundles. 
These  usually  fuse  to  some  extent  so  that  there  come  to  be  only 
about  six  or  eight.  These  remain  easily  distinguishable,  since 
the  primary  medullary  rays  are  very  broad.  The  growth  of 
the  cambium  produces  no  true  phloem  elements  opposite  the  me- 
dullary rays,  although  there  is  some  thin-walled  parenchyma. 

A  crescent-shaped  area  of  stereom  is  finally  formed  at  the 
outer  edge  of  each  phloem  bundle. 

The  pith  becomes  rather  small  in  extent.  There  is  a  perime- 
dullary zone  of  two  or  three  layers  of 
smaller  cells.  According  to  Flot 
[1893]  these  form  at  a  later  time  five 
or  six  layers  of  sclerotic  paren- 
chyma. The  formation  of  cork  was 
not  observed.  It  does  not  take  place 
the  first  year. 

Comparison  of  Structure  of  Hypo- 
cotyl and  Epicotyl, 

The  hypocotyl  has  a  thicker  cor- 
tex than  the  epicotyl :  it  is  without 
a  collenchymatous  hypoderma.  The 
endodermis  is  much  more  distinct  in 
the  former  region  and  the  peculiar  dis- 
tribution of  starch  in  the  pericycle  is 
more  pronounced.  FIG.  6. 


Menisperniuni 
canadeiise 


100  MINNESOTA    BOTANICAL    STUDIES. 

Concerning  the  structure  of  the  stele  it  is  to  be  noted  that  in 
the  hypocotyl  it  is  quadrangular;  it  has  but  four  vascular 
bundles  instead  of  from  eight  to  twelve  and  there  is  no  stereom, 
while  in  the  epicotyl  a  crescentic  mass  of  stereom  borders  each 
phloem  bundle. 

CALYCANTHACE^E. 

Butneria  florida. 
Structure  of  Hypocotyl. 

The  epidermis  consists  of  cells  which  are  nearly  square  in 
cross  section.  They  soon  become  more  or  less  broken,  owing 
to  the  early  formation  of  cork.  A  few  short,  pointed,  uni- 
cellular hairs  are  present. 

A  true  hypoderma  becomes  differentiated  late  in  the  season. 
About  three  or  four  of  the  sub-epidermal  layers  of  cells  become 
collenchymatous.  The  cortex  has  about  twenty  layers  of  cells 
all  approximately  the  same  size.  Intercellular  spaces  abound. 

The  endodermal  cells  are  but  slightly  smaller  than  those  of 
the  cortex.  The  endodermis  remains  more  or  less  distinct  until 
the  close  of  the  first  year.  Starch  grains  are  very  small.  A 
few  are  found  in  the  endodermis,  but  no  starch  is  present  in  the 
other  parts  of  the  hypocotyl  till  late  in  the  season,  when  it  is 
found  in  great  abundance  throughout  all  the  parenchymatous 
tissues. 

The  stele  is  somewhat  quadrangular.  There  are  four  xylem 
bundles  and  four  principal  phloem  bundles.  These  are  situated 
in  the  angles  of  the  stele.  There  are  also  some  small  phloem 
areas.  Their  location  will  be  seen  by  reference  to  the  diagram. 
The  phloem  and  xylem  soon  form  narrow,  closed  zones.  The 
former  is  most  developed  at  the  original  angles  of  the  stele. 

It  is  .stated  by  De  Bary  [1884],  that  in  the  seedlings  of  Caly- 
canthaceae  a  transverse  section  of  the  hypocotyl  shows  six  bun- 
dles. In  the  plant  under  investigation,  the  present  writer  found 
this  to  be  true  only  for  the  upper  end  of  the  hypocotyl  where  the 
cortical  bundles,  to  be  mentioned  later,  are  separating  and  pre- 
paring to  leave  the  stele.  This  appearance  is,  of  course,  only 
seen  after  the  fusion  of  the  primaiy  xylem  bundles  in  pairs, 
and  before  complete  rings  of  phloem  and  xylem  are  produced. 

Toward  the  upper  limit  of  the  hypocotyl  there  is  present  a 
small  stereom  bundle  at  each  of  the  four  angles  of  the  stele. 


Ramaley :   HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.     101 

These  stereom  bundles  bend  outward  and  accompany  the  corti- 
cal bundles  in  succeeding  internodes. 

The  pith  is  rather  thick-walled ;  the  cells  are  about  the  same 
size  as  those  of  the  cortex.  A  small-celled  perimedullary  zone 
of  one  or  two  layers  is  at  length  clearly  distinguishable. 

Cork  formation  begins  very  early  in  the  outermost  sub-epider- 
mal layer  of  the  cells. 

Structure  of  EpicotyL 

The  cells  of  the  epidermis  when  seen  in  cross  section  are  rec- 
tangular with  the  long  diameter  parallel  to  the  surface  of  the 
section.  There  are  numerous  pointed  hairs  of  various  lengths. 

A  collenchymatous  hypoderma,  four  or  five  layers  of  cells  in 
thickness,  forms  the  outer  part  of  the  cortex,  the  rest  of  which  is 
composed  of  very  loose  parenchyma. 

A  definite  endodermis  was  not  distinguished.  The  endoder- 
mal  region  is,  however,  easily  recognized  by  the  presence  of 
starch  in  many  of  the  cells.  Starch  is  afterward  found  in  great 
abundance  in  pith,  cortex  and  medullary  rays. 

The  normal  phloem  and  xylem  form  closed  rings  even  in  the 
youngest  material  examined.  In  the  cortex,  about  half  way 
between  the  epidermis  and  phloem  are  four  vascular  bundles, 
ninety  degrees  apart ;  each  bundle  consists  of  a  more  or  less 
crescent-shaped  mass  of  lignified  sclerenchyma,  at  whose  con- 
cave surface  is  a  small  .area  of  slightly  lignified  xylem,  consist- 
ing usually  of  five  to  ten  cells.  Adjoining  this  xylem  and  pro- 
jecting some  distance  toward  the  stele  is  a  lenticular  mass  of 
phloem.  The  general  arrangement  of  the  bundle  is  the  same 
as  that  carefully  described  for  Calycanthus  sp.  by  Woronin 
[1860]  and  for  Calycanthus  occidentalis  by  Williams  [1894]. 
Serial  sections  showed  that  in  this  species  these  cortical  bundles 
enter  the  stele  about  i  mm.  below  the  insertion  of  the  cotyledons, 
and  not  at  the  middle  of  the  first  internode  as  reported  by  Herail 
[1885]  for  certain  other  species. 

The  pith  is  large.  There  is  a  definite  perimedullary  zone  of 
about  three  layers  of  small  cells. 

The  cork  cambium  is  formed  very  early  in  the  outermost 
hypodermal  layer  (cf.  Moeller  [1882],  p.  364). 

Comparison  of  Structure  of  Hypocotyl  and  EpicotyL 
The  epidermal  hairs  of  the  hypocotyl  are  fewer  and  shorter 


102 


MINNESOTA    BOTANICAL    STUDIES. 


than  those  of  the  epicotyl,     There  is  in  the  former  region  also 
a  less  developed  hypoderma. 

The  stele  of  the  hypocotyl  is 
originally  quadrangular ;  it  is  sur- 
rounded by  a  distinct  endodermis, 
and  has  four  xylem  bundles  and 
four  principal  phloem  bundles.  The 
stele  of  the  epicotyl  is  cylindrical, 
without  a  distinct  endodermis,  and 
even  at  a  very  early  age,  the  xylem 
and  phloem  form  closed  rings. 

The  hypocotyl  has  no  cortical 
vascular  bundles ;  of  these  the  epi- 
cotyl has  four. 

The  presence  of  true  collenchy 
matous  hypoderma  in  the  hypocotyl 
deserves  special  mention,  as  this 
forms  an  exception  to  the  general 
rule  that  collenchyma  is  not  de- 
veloped in  the  hypocotyl. 

C^ESALPINACE^E. 

Parkinsonia  aculeata. 

Structure  of  Hypocotyl. 

The  epidermis  is  composed  of  cells  which  are  rectangular  in 
cross  section.  They  are,  at  first,  radially  elongated.  Eventu- 
ally they  become  elongated  in  the  other  direction. 

The  cortex  is  many-layered.  There  is  no  hypoderma.  The 
outer  cells  of  the  cortex  are  much  smaller  than  those  further 
down.  Very  early  in  the  history  of  the  hypocotyl  a  parenchy- 
matous  sheath  of  small  cells  is  formed  in  the  cortex  about  mid- 
way between  epidermis  and  endodermis.  The  cells  are  not 
arranged  in  definite  rows.  The  position  of  this  sheath  is  shown 
in  the  last  plate  accompanying  this  paper. 

The  cells  of  the  endodermis  are  smaller  than  those  of  the  cor- 
tex. They  contain  starch.  The  endodermis  is  quite  distinct; 
it  was,  however,  not  definitely  distinguished  in  material  col- 
lected late  in  the  season.  The  cortex  and  pith  at  a  later  time 
also  have  some  starch. 

The  stele  is  four-angled.     There  are  originally  four  phloem 


RamaJcy :   HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.     103 


bundles  and  eight  paired  xylem  bundles.  Eventually  closed 
rings  of  xylem  and  phloem  are  formed. 

At  the  corners  of  the  stele  in  the  pericycle  groups  of  scleren- 
chyma  are  formed.  The  cells  become  very  thick-walled  and 
each  group  quite  large. 

The  pith  is  large.  The  cells  are  rather  thick-walled.  No 
definite  perimedullary  zone  was  distinguished. 

In  the  material  examined  cork  formation  had  not  commenced. 

Structure  of  EpicotyL 

The  epidermal  cells  are  thin-walled,  square  in  cross  section, 
becoming  at  length  much  flattened.  There  is  no  collenchyma. 
The  cortex  is  narrow ;  the  cells  are  about  the  same  size  as  those 
of  the  epidermis. 

The  endodermis  is  composed  of  thin-walled  cells.  After 
secondary  growth  of  the  stelar  tissues  it  cannot  be  definitely 
seen.  The  cells  are  about  the  same  size  as  those  of  the  cortex ; 
they  contain  starch.  Starch  is  found  at  a  later  time  in  the 
various  parenchymatous  tissues. 

The  number  of  vascular  bundles  is  variable.  Usually  there 
are  about  twelve.  These,  at  length,  fuse  to  form  closed  rings 
of  phloem  and  xylem. 

The  pericycle  develops  a  sheath  of  sclerenchyma  which  almost 
completely  shuts  in  the  phloem.  The  cells  were  not  very  thick- 
walled  in  the  material  examined. 

The  pith  is  large,  the  cells  rather 
thin-walled.  A  perimedullary  zone 
of  small-celled  parenchyma  at  length 
becomes  differentiated. 

No  material  old  enough  to  show 
cork  formation  was  examined. 


Comparison  of  Structure  of  Hypo- 
cotyl  and  EpicotyL 

The  hypocotyl  differs  from  the  epi- 
cotyl  in  having  a  thicker  cortex  with 
a  narrow  small-celled  parenchyma- 
tous sheath.  The  cells  of  the  cortex 
are  also  larger. 

In  its  primary  stelar  structure  the 
differences  are  very  marked.  The 


1'nrkiiiMMiia 
aculenta 


FIG.  8. 


104  MINNESOTA    BOTANICAL    STUDIES. 

hypocotyl  has  four  phloem  bundles  and  eight  xylem  bundles 
instead  of  a  large  number  of  conjoint  bundles.  It  has  four 
large  groups  of  stereom  instead  of  a  narrow,  almost  continuous 
sclerenchymatous  sheath. 

Cercis  canadensis. 

Structure  of  Hypocotyl. 

The  epidermal  cells  are  rectangular  in  cross  section ;  the 
radial  diameter  is  the  longer.  These  cells  never  become  tan- 
gentially  elongated.  The  cells  of  the  outer  cortex  are  smaller 
than  those  within.  An  indefinite  sheath  of  small-celled  paren- 
chyma similar  to  that  in  Parkinsonia  can  sometimes  be  recog- 
nized. 

The  endodermis  is  small-celled ;  it  remains  distinct  through 
the  first  year.  Its  cells  contain  starch.  Starch  is  also  present 
toward  the  close  of  the  year  in  the  pith. 

The  stele  is  originally  quadrangular.  There  are  four  xylem 
bundles  and  four  phloem  bundles.  These,  at  length,  develop 
into  closed  rings. 

Four  small  groups  of  sclerenchyma  make  their  appearance 
in  the  pericycle  at  an  early  time  and  become,  at  length,  consid- 
erably extended. 

The  pith  finally  becomes  cylindrical.  The  cells  are  large 
and  thin-walled.  The  perimedullary  zone  is  not  clearly  dif- 
ferentiated. 

Cork  formation  takes  place  in  the  cortex  either  next  to  or 
very  near  the  endodermis.  It  begins  sometime  before  the  close 
of  the  season. 

Structure  of  Epicotyl. 

The  epicotyl  is  somewhat  quadrangular  in  the  early  stages. 
The  epidermal  cells  are  rectangular  in  cross  section.  The 
tangential  diameter  is  the  greater.  There  is  no  hypoderma. 
The  cortex  is  thin.  The  cells  are  all  about  the  same  size. 

The  endodermis  was  not  definitely  distinguished.  In  the 
youngest  material  examined  the  phloem  forms  a  closed  ring. 
There  are  four  large  primary  xylem  bundles.  There  are  also 
some  smaller  ones.  The  latter  have  often  only  one  or  two 
xylem  cells.  A  closed  zone  of  xylem  is  soon  produced. 

Nearly  all  the  cells  of  the  pericycle  become,  at  length,  scler- 
otic, thus  forming  an  almost  continuous  sheath  with  but  few 
parenchymatous  cells. 


Ramaley :   IIYPOCOTYL  AND  EPICOTVL  IN  WOODY  PLANTS.     105 


The  pith  is  large-called.  A  perimedullary  zone  was  not  dis- 
tinguished. 

Cork  formation  takes  place  in  the  second  cortical  layer  as  in 
Cercis  siliquastrum  (fide  Moeller  [1882]  ). 

Comparison  of  Structure  of  Hypocotyl  and  Epicolyl. 

The  epidermal  cells  of  the  hypocotyl,  when  seen  in  cross 
section,  appear  radially,  not  tan- 
gentially  elongated  as  in  the  epi- 
cotyl.  In  the  former  region  the 
endodermis  is  distinct,  the  cortex 
thicker  and  the  sclerenchyma  at  first 
differently  disposed. 

The  stele  of  the  hypocotyl  has 
originally  four  phloem  bundles  and 
four  xylem  bundles.  The  young- 
est material  of  the  epicotyl  which 
was  examined  has  a  closed  ring 
of  phloem  and  four  large  xylem 
bundles,  also  a  few  small  groups  of 
xylem. 

Cork  formation  in  the  hypocotyl 
takes  place  in  the  lower  cortex ; 
in  the  epicotyl  it  takes  place  in  the 
second  cell  layer  of  the  cortex. 


Cercis  canadensis 
FlG.  9. 


Gleditsia  triacanthos. 
Structure  of  Hypocotyl. 

The  epidermis  is  composed  of  rather  thick-walled  cells  which 
are  oblong  in  cross  section,  the  long  axis  being  at  right  angles 
to  the  periphery  of  the  section.  These  cells  are  eventually 
elongated  in  the  tangential  direction. 

The  cortex  is  very  thick.  There  is  no  differentiated  hypo- 
derma,  but  three  or  four  of  the  outer  cortical  layers  are  com- 
posed of  smaller  cells  than  those  below. 

The  endodermis  is  definite ;  it  is  large-celled.  In  some 
places  it  is  two  layers  of  cells  in  thickness.  Starch,  at  first 
present  only  in  the  endodermis,  is  eventually  widely  distributed 
throughout  all  the  parenchymatous  tissues. 

The  stele  is  cylindrical.     There  are  in  the  young  hypocotyl 


106  MINNESOTA    BOTANICAL    STUDIES. 

eight  paired  xylem  bundles  and  a  large  number  of  groups  of 
phloem.  The  latter  soon  grow  together,  forming  a  complete 
ring,  while  the  xylem  bundles  first  fuse  in  pairs,  afterward 
growing  together  into  a  closed  zone. 

In  the  pericycle,  alternating  with  the  paired  xylem  bundles 
there  are  developed  four  large  bands  of  sclerenchyma  which  ex- 
tend so  far  around  that  they  nearly  touch  each  other.  By  the 
end  of  the  first  season  these  become  divided  into  a  number  of 
groups  by  the  intercalation  of  parenchymatous  cells. 

The  pith,  which  is  eventually  of  slight  extent,  is  composed  of 
large-celled  parenchyma. 

Cork  formation  begins  rather  early  the  first  season  in  the 
third  or  fourth  cell  layer  of  the  cortex. 

Structure  of  Epicotyl. 

The  general  shape  of  the  epicotyl  is  originally  somewhat 
hexagonally  prismatic ;  it  soon  becomes  cylindrical. 

The  cells  of  the  epidermis  are  originally  nearly  square  in 
cross  section.  There  are  numerous,  long,  curved,  pointed  epi- 
dermal hairs.  The  outer  two  layers  of  the  cortex  become 
slightly  collenchymatous.  The  other  cortical  layers  are  com- 
posed of  parenchyma. 

The  endodermis  was  not  distinguished  in  material  taken  in 
the  autumn  but  in  the  young  epicotyl  is  quite  distinct.  The  cells 
are  rather  large,  similar  to  those  of  the  cortical  region  but  packed 
with  starch. 

The  phloem,  in  youngest  material  examined,  forms  a  ring  of 
tissue.  There  are  about  six  principal  xylem  bundles  which 
soon  fuse. 

A  broken  sclerenchymatous  ring  is  formed  which  resembles 
that  of  the  epicotyl.  No  other  stereom  is,  as  a  rule,  produced 
the  first  year. 

The  pith  is  large  and  composed  of  cells  with  unlignified  walls. 
There  is  a  small-celled  perimedullary  zone. 

Cork  formation  takes  place  in  the  hypoderma  (cf.  Moeller 
[1882],  p.  393). 

Comparison  of  Structure  of  Hypocotyl  and  Epicotyl. 

The  hypocotyl  differs  from  the  epicotyl  in  the  absence  of  epi- 
dermal hairs  and  of  a  collenchymatous  hypoderma,  in  the  pri- 
mary structure  of  the  stele,  and  in  its  smaller  pith. 


Ramaley :   IIYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.     107 


In  the  hypocotyl  there  are  at  first 
four  pairs  of  xylem  bundles  and  a 
number  of  phloem  bundles.  Four 
large  groups  of  sclerenchyma  soon 
make  their  appearance  in  the  peri- 
cycle.  In  the  epicotyl,  on  the 
other  hand,  a  closed  ring  of  phloem 
surrounds  usually  about  six  xylem 
bundles.  No  differentiated  peri- 
medullary  zone  was  distinguished 
in  the  hypocotyl. 

The  structure  of  the  two  regions 
at  the  close  of  the  first  year  differs 
only  in  the  perimedullary  region  and 
pith  ;  the  formation  of  cork  having 
removed  the  epidermis  and  hypo- 
derma. 


Gleditsia 
triacanthos 


FIG.  10. 


PAPILIONACE^E. 

Amorpha  fruticosa. 

Structure  of  Hypocotyl. 

The  epidermis  consists  of  cells  rather  small,  somewhat  thick- 
walled,  square  or  nearly  so,  in  cross  section,  at  length  becom- 
ing flattened.  The  cells  of  the  cortex  are  large ;  those  imme- 
diately below  the  epidermis  somewhat  smaller,  but  not  forming 
a  definite  hypoderma. 

The  endodermis  of  thin-walled  cells  containing  starch  re- 
mains distinct  for  some  time.  Toward  the  close  of  the  first  year 
its  exact  position  cannot  be  determined,  although  it  can  be  lo- 
cated approximately.  A  small  amount  of  starch  is  scattered 
throughout  the  cortex,  pith  and  pericycle  as  well  as  the  endo- 
dermis, even  in  the  youngest  stage.  This  is  not  the  case  in 
most  species.  Later  the  phloem  and  the  medullary  rays  also 
come  to  be  filled  with  starch. 

The  stele  is  at  first  quadrangular,  and  remains  so  for  a  con- 
siderable length  of  time.  There  are  four  conjoint  vascular 
bundles,  and  in  addition  there  appear  a  few  small  patches  of 
phloem.  The  bundles  soon  tend  to  unite  in  pairs.  Xylem  and 
phloem  at  length  form  complete  zones.  The  medullary  rays 
are  very  numerous ;  they  are  one  cell  in  width. 


108  MINNESOTA    BOTANICAL    STUDIES. 

About  the  time  that  the  epicotyl  has  reached  its  full  length 
four  small  areas  of  sclerenchyma  appear  in  the  pericycle,  one 
adjoining  the  phloem  of  each  vascular  bundle.  These  increase 
somewhat  in  size,  and  are  still  visible  in  two-year-old  material. 
Numerous  isolated  sclerenchymatous  elements  are  found  scat- 
tered through  the  phloem. 

The  pith  is  composed  of  rather  large  cells  with  thin  walls, 
which  soon  become  lignified.  As  the  plant  grows  older  the  pith 
becomes  almost  obliterated.  No  perimedullary  zone  was  dis- 
tinguished. 

Cork  formation  takes  place  in  the  outer  pericycle,  at  length 
cutting  off  all  tissues  outside,  leaving  the  bundles  of  scleren- 
chyma which  are  at  the  inner  limit  of  the  pericycle. 

Structure  of  Epicotyl. 

The  epidermis  is  composed  of  cells  nearly  square  in  cross 
section.  These  abut  directly  upon  a  large-celled,  few-layered 
cortex.  There  is  no  hypoderma. 

A  definite  endodermis  was  not  distinguished  at  any  time  al- 
though in  a  very  young  stage  certain  starch  containing  cells 
were  recognized  as  having  the  appearance  of  endodermis ;  a 
continuous  ring  of  them  was  not  traced.  With  the  exception  of 
the  endodermal  and  medullary  region,  starch  does  not  occur 
until  the  plant  has  developed  a  number  of  internodes  above  the 
epicotyl.  The  cells  of  pericycle  and  phloem  are  at  length  filled 
with  starch. 

The  stele  is  cylindrical  from  the  first.  Owing  to  fusions  the 
number  of  vascular  bundles  is  variable.  There  are,  however, 
generally  about  five  or  six  bundles.  The  phloem  and  xylem 
eventually  form  closed  rings. 

There  is  a  narrow  interrupted  ring  of  stereom  at  the  outer 
edge  of  the  pericycle.  Toward  the  end  of  the  first  season 
numerous  small  patches  of  thick-walled  fibers  appear  in  the 
phloem  and  the  pericycle. 

The  pith  is  large-celled ;  it  does  not  decrease  appreciably  in 
size  as  the  stem  grows  older. 

In  the  lower  part  of  the  epicotyl  cork  formation  takes  place 
in  the  pericycle  below  the  ring  of  stereom  mentioned  above, 
thus  cutting  off  the  cortex  and  epidermis  which  soon  die  and 
disappear.  In  the  upper  part  it  takes  place  in  the  cortex 
(cf.  Moeller  [1882],  p.  383).  This  plant  shows  a  distinct  "  re- 
gion tigellaire  "  in  two-year-old  material. 


Ramaley :   HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.     109 

Comparison  of  Structure  of  Hypocotyl  and  Epicotyl. 

In  very  young  plants  the  hypocotyl  shows  a  few  slightly  dif- 
ferentiated layers  of  smaller  cells  in  the  outer  cortex.  The 
epicotyl  has  nothing  of  the  kind. 

The  endodermis  of  the  hypocotyl 
is  distinct  in  the  early  stages,  but  was 
not  definitely  located  in  the  epicotyl. 
Starch  appears  earlier  in  the  hypocotyl 
and  the  four  large  groups  of  stereom 
are  not  represented  at  all  in  the  epi- 
cotyl which,  however,  has  an  inter- 
rupted circle  of  the  same  material. 

In  the  former  region  the  stele  is  at 
first  quadrangular,  while  always  cylin- 
drical in  the  latter.  Cork  formation 
in  the  hypocotyl  is  pericyclic,  but  is 
cortical  in  the  epicotyl. 

The  final  structure  of  the  two  regions 
is  very  similar,  save  in  the  arrange- 
ment of  sclerenchyma. 


Amorpha 

fruticosa 
FIG.   n. 


Robinia  pseudacacia. 
Structure  of  Hypocotyl. 

The  epidermal  cells  are  oblong  in  cross  section,  radially 
elongated  at  first,  later  becoming  elongated  in  the  tangential 
direction.  A  few  straight  multicellular  hairs  are  present. 
There  is  no  differentiated  hypoderma.  The  cells  of  the  cortex 
are  all  about  the  same  size. 

The  endodermis  is  small-celled  and  contains  starch.  It  is  not 
easily  distinguished  in  material  taken  at  the  close  of  the  grow- 
ing season.  Starch  is  also  found  in  some  of  the  pericyclic  cells 
in  early  stages.  Later  nearly  all  the  parenchymatous  tissues 
have  starch. 

The  stele  is  originally  quadrangular.  There  are  eight  phloem 
bundles  and  four  xylem  bundles.  These  soon  produce  closed 
zones. 

In  the  pericycle  opposite  each  of  the  original  xylem  bundles 
a  group  of  stereom  appears.  These  groups,  at  length,  become 
quite  large.  In  addition  to  these,  at  the  close  of  the  first  year, 
there  are  some  small  patches  of  stereom  irregularly  disposed 
just  outside  the  phloem. 


110 


MINNESOTA    BOTANICAL    STUDIES. 


The  pith  is  composed  of  parenchymatous  cells  which  acquire 
thick  lignified  walls.  There  is  a  well-differentiated  perimedul- 
lary  zone  three  or  four  cells  in  width.  The  cells  are  small  and 
have  thick  lignified  walls. 

Cork  formation,  according  to  Flot  [1890],  takes  place  rather 
deep  in  the  cortex. 

Structure  of  Epicotyl. 

The  cells  of  the  epidermis  are  square  or  oblong  in  cross  sec- 
tion, and  become  in  time  greatly  flattened.  There  are  numerous 
epidermal  hairs.  A  narrow  collenchymatous  hypoderma  is 
present.  The  cells  of  the  cortex  are  about  the  same  size  as  the 
epidermal  cells. 

The  endodermis  was  distinguished  only  in  very  young  stages. 
The  cells  are  rather  small  and  closely  packed  with  starch. 
Starch  is  found  at  a  later  time  in  the  various  parenchymatous 
tissues.  Troschel  [1879]  states  that  in  year-old  twigs  starch  is 
present  in  some  of  the  elements  of  the  wood  but  disappears  the 
next  year. 

In  young  material  the  epicotyl  is  elliptical  in  cross  section. 
The  stele  follows  this  closely  in  shape.  The  phloem  forms  a 
closed  ring  surrounding  a  variable  number  of  xylem  bundles. 
There  are  usually  more  than  eight  of  these  bundles.  The 
xylem  also  soon  forms  a  complete  zone  in  which  medullary  rays 

are  prominent. 

The  pith  cells  become,  at  length, 
thick-walled.  There  is  a  well-de- 
fined perimedullary  zone. 

Cork  arises  in  the  fourth,  fifth  or 
sixth  layer  of  the  cortex  (cf.  Moel- 
ler  [1882],  p.  384). 

Comparison  of  Structure  of  Hypo- 

cotyl  and  Epicotyl. 
The  hypocotyl  is  without  the  col- 
lenchymatous hypoderma  of  the  epi- 
cotyl ;  it  has  fewer  epidermal  hairs ; 
the  cortex  is  thicker ;  there  are  four 
large  groups  of    stereom  with  some 
very  small  ones  instead  of  a  broken 
FIG.  12.  ring  °f  medium-sized  bundles. 


Robin  ia 

pseudacacia 


Ramalcy  :   HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.     Ill 

In  its  primary  structure  the  stele  of  the  hypocotyl  differs  con- 
siderably from  that  of  the  epicotyl.  There  are  four  xylem 
bundles  instead  of  eight  or  more  and  eight  phloem  bundles  in- 
stead of  a  closed  ring  of  phloem. 

The  cork,  although  of  cortical  origin  in  both  regions,  arises 
in  the  hypocotyl  in  deeper  layers. 


Ptelea  trifoliata. 
Structure  of  Hypocotyl. 

The  cells  of  the  epidermis,  when  seen  in  cross  section,  appear 
nearly  square.  They,  at  length,  are  flattened.  There  are  a 
few  short,  blunt,  unicellular  hairs.  The  cortex  is  large-celled. 
There  is  no  distinct  hypoderma  differentiated. 

The  endodermis  is  large-celled  and  contains,  at  first,  very 
little  starch.  It  later  becomes  closely  packed  with  starch.  The 
various  parenchymatous  tissues  at  length  also  contain  starch 
in  the  cell  cavities.  Numerous  lysigenous  reservoirs  are  pres- 
ent in  the  outer  part  of  the  primary  cortex. 

The  stele  is  originally  four-angled.  There  is  one  phloem 
bundle  and  one  xylem  bundle  in  each  angle.  The  phloem  soon 
forms  a  closed  ring  surrounding  the  now  greatly  enlarged  xylem 
bundles  which  enclose,  at  this  stage,  a  somewhat  cruciform  pith. 
The  xylem  bundles  also  finally  fuse. 

Four  very  small  groups  of  sclerenchyma  appear,  toward  the 
end  of  the  season,  in  the  pericycle.  They  are  equidistant. 
There  are  about  six  cells  in  each  group.  Some  sections  do  not 
show  all  these  groups,  as  the  sclerenchymatous  elements  do  not 
form  continuous  strands  in  the  hypocotyl.  Some  sections  show 
no  sclerenchyma  at  all. 

The  pith  is  eventually  quite  small.  The  perimedullary  zone 
is  not  well  developed. 

The  formation  of  cork  begins  early  in  the  outermost  cortical 
layer  of  cells. 

Structure  of  EpicotyL 

The  epidermis  is  composed  of  cells  which  appear  slightly  rec- 
tangular in  cross  section.  They  are  elongated  in  the  tangen- 
tial direction.  Numerous  epidermal  hairs  are  present.  There 
is  a  narrow  collenchymatous  hypoderma.  The  cells  of  the  in- 
ner cortex  are  very  large. 


112 


MINNESOTA    BOTANICAL    STUDIES. 


The  endodermis  is  distinct  and  can  be  recognized  in  year-old 
material.  Starch  is  present  from  the  first.  The  various  paren- 
chymatous  tissues  at  length  have  a  small  amount  of  starch. 
Secretion  cavities  develop  in  the  cortex. 

The  stele  is  small ;  much  smaller  than  is  usual  in  most  spe- 
cies. In  the  youngest  material  examined  the  phloem  forms  a 
complete  ring  surrounding  a  small  number  of  xylem  bundles 
which  eventually  fuse. 

Numerous  groups  of  elements  in  the  pericycle  become  scler- 
otic so  that  they  form  an  interrupted  ring  of  sclerenchyma  sur- 
rounding the  phloem. 

The  pith  is  small,  unusually  so  for  an  epicotyl.  There  is  a 
definite  perimedullary  zone  of  small-celled  parenchyma  contain- 
ing starch. 

Thepiormation  of  cork  takes  place  in  the  outermost  layer  of 
hypoderma  (cf.  Moeller  [1882],  p.  326). 

Comparison  of  Structure  of  Hypocotyl  and  Epicotyl. 

The  hiypocotyl  does  not  have  the  epidermal  hairs  and  the  collen- 

chymatous  hypoderma  of  the  epicotyl. 
The  cortex  of  the  former  region,  though 
very  thick,  is  but  little  thicker,  in  pro- 
portion, than  that  of  the  epicotyl. 

In  the  stele  of  the  hypocotyl  there 
are  four  phloem  bundles  and  an  equal 
number  of  xylem  bundles,  while  in  the 
epicotyl,  in  the  youngest  material  ex- 
amined, the  phloem  forms  a  closed  ring 
surrounding  about  six  xylem  strands. 

The  sclerenchyma  of  the  hypocotyl 
is  in  four  somewhat  irregular  columns 
in  the  pericycle,  while  in  the  epicotyl 
it  forms  more  nearly  a  closed  sheath. 
In  the  former  region  also  the  peri- 
medullary  zone  is  poorly  developed. 


Ptelea  trifoliate 


FIG.  13. 

SIMARUBACE^. 

Ailanthus  glandulosa. 

Structure  of  Hypocotyl. 

The  epidermis  consists  of  small  cells,  square  or  nearly  so,  in 
cross  section,   and  considerably  bulged   when  young.     A  few 


Ramaley :   HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.     113 

short,  unicellular  hairs  were  seen,  but  none  noted  in  very  young 
plants.  There  is  a  hypoderma  of  one  or  two  layers  of  some- 
what larger  and  thicker-walled  cells.  The  rest  of  the  cortex  is 
parenchymatous  and  large-celled. 

The  endodermis  consists  of  cells  somewhat  smaller  than  those 
of  the  adjacent  cortical  layer.  The  endodermis  was  not  recog- 
nized in  older  material.  Starch  is  present  from  the  first  in  the 
endodermis,  pericycle  and  pith,  but  does  not  appear  in  the  cor- 
tex till  nearly  the  close  of  the  first  season. 

The  stele  is  at  first  quite  small.  There  are  four  xylem  bun- 
dles arranged  in  pairs  and  four  phloem  bundles  similarly  dis- 
posed. The  phloem  soon  forms  a  complete  ring,  surrounding 
the  now  considerably  enlarged  xylem  bundles,  which  also  even- 
tually form  a  closed  ring. 

Opposite  each  of  the  four  original  xylem  bundles  there  ap- 
pears in  the  pericycle  a  group  of  sclerenchymatous  cells. 
These  groups  become,  at  length,  somewhat  divided  so  that  the 
old  hypocotyl  may  have  a  considerable  number  of  smaller 
groups.  There  are  numerous  sclerenchymatous  fibers  scattered 
in  small  and  large  patches  through  the  phloem  and  pericycle. 

The  pith  is  thin-walled ;  toward  the  end  of  the  first  season  it 
becomes  lignified.  The  perimedullary  zone,  described  by  Flot 
[1893],  as  an  important  feature  of  the  stem  structure  is  first 
definitely  noted  at  this  time. 

The  oleoresin  canals  described  by  Trecul  [1867]  as  occurring 
at  the  outer  border  of  the  pith,  and  by  Van  Tieghern  [1884]  as 
in  the  inner  xylem  of  the  stem,  were  not  distinguished  in  the 
hypocotyl.  Crystal  rosettes  of  calcium  oxalate  occur  singly  in 
certain  cells  of  the  phloem  area.  Single  oleoresin  cells  are 
found  here  and  there  in  the  cortex  and  phloem. 

Cork  formation,  as  noted  by  Flot  [1889  and  1890]  takes 
place  in  the  layer  of  cells  just  below  the  epidermis. 

Structure  of  Epicotyl. 

The  epidermis  resembles  that  of  the  hypocotyl,  but  there  are 
numerous,  somewhat  long,  curved  or  hooked  epidermal  hairs. 
Most  of  these  are  unicellular. 

The  hypoderma  is,  as  previously  described  for  the  stem  by 
De  Bary  ([1884],  p.  404),  collenchymatous.  The  cells  are 
small ;  toward  the  inside  the  hypoderma  gradually  shades  into 
the  ordinary  cortex. 


114 


MINNESOTA    BOTANICAL    STUDIES. 


A  definite  endodermis  was  not  distinguished  at  any  stage,  al- 
though, since  starch  is  present  in  the  region  of  the  pericyle  and 
endodermis  from  the  first,  those  regions  can  be  located  approx- 
imately. Starch  is  found  later  in  pith  and  cortex  ;  also  in  many 
of  the  inner  xylem  elements. 

There  are,  at  first,  eight  to  ten  conjoint  vascular  bundles. 
Eventually  the  phloem  and  xylem  form  closed  rings. 

Scattered  sclerenchymatous  elements  are  found  in  the  phloem, 
pericycle  and  cortex. 

The  pith  is  irregular  in  outline.  The  first  formed  xylem  ele- 
ments project  into  it.  The  perimedullary  zone  is  not  conspic- 
uous the  first  year,  being  composed  of  a  few  cells  with  unligni- 
fied  walls. 

The  cork  cambium  is  formed  in  the  outermost  hypodermal 
layer  (cf.  Moeller  [1882],  p.  327). 

Comparison  of  Structure  of  Hypocotyl  and  Epicotyl. 

The  hypocotyl  has  a  few,  the  epi- 
cotyl  a  considerable  number,  of  epi- 
dermal hairs.  The  hypocotyl  does 
not  have  the  collenchymatous  hypo- 
derma  found  in  the  epicotyl.  The 
pith  is  smaller  and  circular  instead 
of  scalloped  ;  the  perimedullary  zone 
is  better  developed. 

The  endodermis  is  distinct  in  the 
hypocotyl  for  a  considerable  time, 
while  in  the  epicotyl  it  was  not 
definitely  distinguished  at  all.  The 
hypocotyl  has,  at  first,  four  xylem 
and  four  phloem  bundles  ;  the  epi- 
cotyl eight  to  ten  conjoint  bundles. 

At  the  close  of  the  year  the  only  dif- 
ferences are  those  noted  in  the  me- 
dullary and  perimedullary  regions. 


Ailanthiis 
glandulosa 


FIG.  14. 

ANACARDIACE^E. 

Schinus  molle. 
Structure  of  Hypocotyl. 

The  epidermal  cells  are  square  or  oblong  in  cross  section, 
becoming,  at  length,  flattened.     There  are  numerous  short  epi- 


Ramaley :   HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.     115 

dermal  hairs.  No  hypoderma  is  developed  but  the  cells  of 
three  or  four  outer  layers  of  the  cortex  are  smaller  than  those 
of  deeper  layers. 

The  endodermis  is  small-celled  and  easily  recognized  in 
young  stages,  although  at  that  time  the  cells  are  without  starch. 
Later  starch  appears  in  small  quantities  in  these  cells  and  in 
those  of  the  pith  and  phloem. 

The  stele  is  originally  four-angled  and  remains  so  for  some 
time.  In  each  angle  there  is  a  single  xylem  bundle  and  two 
groups  of  phloem ;  these  form  a  crescent-shaped  mass  border- 
ing a  group  of  cells  which  later  develop  into  a  resin  duct. 
After  a  time  secondary  vascular  bundles  are  intercalated  be- 
tween the  primary  bundles.  All  finally  fuse  to  produce  closed 
zones  of  phloem  and  xylem. 

A  few  small  groups  of  sclerenchymatous  cells  develop  at  the 
outer  border  of  the  phloem. 

The  pith  remains  somewhat  four-sided.  The  four  original 
xylem  bundles  project  into  it  at  the  angles.  The  pith  cells 
have  thin,  unlignified  walls.  A  perimedullary  zone  of  small 
cells  was  distinguished. 

Material  old  enough  to  show  cork  formation  was  not  obtained. 

Structure  of  Epicotyl. 

The  epidermis  resembles  that  of  the  hypocotyl.  Trichome 
structures  seem  to  be  no  more  abundant.  There  is  no  collen- 
chymatous  hypoderma  developed.  The  cells  of  the  cortex  are 
all  about  the  same  size. 

The  endodermis  is  not  easily  recognized  owing  to  the  fact 
that  in  young  stages  it  contains  no  starch.  Later  when  starch 
is  present  the  cells  have  been  compressed  and  displaced  by 
pressure  from  the  subjacent  tissues. 

The  stele  contains  a  variable  number  of  vascular  bundles. 
Usually  there  are  about  eight.  In  connection  with  each  bundle 
is  a  small  resin  passage,  at  first  pointed  out  by  Trecul  [1867]. 
In  older  material  these  resin  passages  become  quite  large  and 
somewhat  flattened.  The  phloem  and  xylem  then  form  closed 
zones. 

Groups  of  sclerenchyma,  usually  consisting  of  only  a  few 
cells,  are  found  at  the  periphery  of  the  phloem.  These  are 
often  located  near  the  resin  passages. 

The  pith  is  nearly^circular,  not  quadrangular,  and  is  com- 


116 


MINNESOTA    BOTANICAL    STUDIES. 


posed  of  large,  thin-walled  cells.     There  is  a  distinct  perime- 
dullary  zone. 

The  region  of  cork  formation  was  not  determined. 

Comparison  of  Structure  of  Hypocotyl  and  EpicotyL 

In  their  primary  structure  the  steles  of  the  hypocotyl  and  epi- 
cotyl  show  important  differences.  That  of  the  former  region  is 
quadrangular ;  it  has  four  primary  vascular  bundles  and  at  a 

later  time  other  secondary  bundles 
are  intercalated.  These  latter  do  not 
have  resin  canals.  In  the  epicotyl 
there  are  about  eight  vascular  bundles 
each  with  a  resin  canal. 

The  pith  of  the  hypocotyl  is  four- 
sided,  that  of  the  epicotyl  circular  in 
outline,  when  seen  in  cross  section. 

RHAMNACE^E. 
Berchemia  racemosa. 

Structure   of  Hypocotyl. 

The  cells  of  the  epidermis  are  nearly 
square  in  cross  section,  sometimes 
radially  elongated,  but  becoming  at 
length  considerably  flattened.  No 
hypoderma  is  developed,  although  the 
cells  of  the  outermost  layer  of  cortical  tissue  are  considerably 
smaller  than  those  below.  There  are  about  five  layers  of  cells 
in  the  cortex.  This  tissue  is  extremely  loose,  having  many  in- 
tercellular spaces. 

The  endodermis  is  quite  distinct  until  nearly  the  time  that 
cork  formation  begins.  The  cells  are  smaller  than  those  of  the 
cortex  but  larger  than  the  pericyclic  elements. 

Starch  is  present  in  the  endodermis  from  the  first,  but  does 
not  appear  in  the  cortex  at  all,  nor  in  the  pith  and  phloem  till 
about  the  close  of  the  season. 

The  stele  is  originally  four-angled.  There  are  four  xylem 
and  four  phloem  bundles.  These  are  paired.  They  soon  fuse 
so  that  there  are  two  crescent-shaped  bundles,  and  by  further 
growth  closed  rings  of  xylem  and  phloem  are  produced. 

While  the  bundles  are  in  the  crescent  form  four  small  groups 


Schtniil  inolle 


FIG.  15. 


Ramaley :   HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.     117 

of  sclerenchyma  appear  in  the  pericycle,  one  opposite  each  of 
the  original  xylem  groups. 

The  pith  is  composed  of  large  cells,  whose  thin  walls  be- 
come, at  length,  somewhat  lignified.  A  perimedullary  zone  of 
about  two  layers  may  be  distinguished  but  is  not  always  con- 
tinuous the  whole  way  around  the  pith. 

The  cork  has  its  origin  in  the  inner  cortex  or  in  the  endoder- 
mis.  Some  of  the  layers  of  cork  carry  a  brown  pigment. 

Structure  of  EpicotyL 

The  epidermis,  composed  originally  of  small  cells,  square  or 
pentagonal  in  cross  section,  eventually  becomes  strongly  cuticu- 
larized  and  the  separate  elements  very  much  flattened. 

There  is  no  hypoderma  developed.  The  cortex  is  rather 
large-celled,  but  very  narrow,  being  only  three  or  four  layers  of 
cells  in  thickness.  During  the  second  year  the  walls  of  these 
cells  become  conspicuously  pitted.  Many  large  crystals,  chiefly 
cubical  in  form,  are  found  in  this  region. 

The  endodermis,  composed  of  flat  cells  containing  starch  is 
distinct  till  near  the  close  of  the  first  season.  Except  in  the  en- 
dodermis starch  is  absent  until  about  the  end  of  the  first  year's 
growth,  when  it  appears  in  the  pith  and  medullary  rays. 

Even  in  very  young  stages  the  phloem  forms  a  closed  zone 
surrounding  a  ring  of  from  six  to  ten,  but  generally  about  eight, 
xylem  bundles.  These  soon  become  fused.  A  narrow  band 
of  sclerenchyma,  for  the  most  part  only  one  cell  wide,  is  found 
at  the  outer  limit  of  the  phloem ;  it  does  not  form  a  closed  ring, 
but  is  more  or  less  irregular  and  broken.  Small  patches  of 
sclerenchyma  are  found  in  the  phloem  of  two-year-old  seed- 
lings. 

The  pith  is  large-celled ;  the  walls  are  thin  but  slightly  ligni- 
fied. No  perimedullary  zone  was  distinguished. 

The  region  of  cork  formation  was  not  distinguished.  Two- 
year-old  material  was  examined,  but  the  cork  cambium  had  not 
begun  to  form. 

Comparison  of  the  Structure  of  Hypocotyl  and  Epicotyl. 

The  epidermis  of  the  hypocotyl  remains  thinner-walled  and 
exhibits  less  cuticularization  than  that  of  the  epicotyl.  This  is 
to  be  expected,  since  in  the  former  region  cork  is  produced  the 
first  year,  while  in  the  latter  not  till  a  later  period. 


118 


MINNESOTA    BOTANICAL    STUDIES. 


The  hypocotyl  has  at  first  four  xylem  and  four  phloem  bun- 
dles,  the  epicotyl   a   closed  ring  of  phloem   and   about   eight 

xylem  bundles.  In  the  former  area 
there  are  but  four  groups  of  scleren- 
chyma,  while  in  the  latter  there  is  an 
interrupted  circle  of  thick-walled  ele- 
ments just  outside  the  phloem. 

The  pith  of  the  hypocotyl  is  smaller 
than  that  of  the  epicotyl. 


Bcrchemia 
racemosa 


FIG.  16. 


Rhamnus  purshiana. 

Structure  of  Hypocotyl. 

The  cells  of  the  epidermis  are  origi- 
nally square  or  pentagonal  in  cross 
section.  They  become,  at  length,  much 
flattened.  The  cortex  is  thick  and 
very  large-celled.  No  hypoderma  is 
differentiated. 

The  endodermis  consists  of  small,  thin-walled  cells  contain- 
ing starch.  It  remains  distinct  until  cork  formation  takes  place. 
Starch  is  found,  late  in  the  season,  in  the  perimedullary  zone 
and  phloem,  but  not  in  the  cortex. 

The  stele  is  originally  somewhat  four  sided  and  has  four  vas- 
cular bundles  which  soon  fuse  and  produce  closed  zones. 

The  pith  is  composed  of  very  large  cells.  The  perimedul- 
lary zone  is  rather  ill-defined.  It  is  one  or  two  cells  in  width. 
The  cells  are  small  and  contain  starch. 

The  cork  is  of  endodermal  origin.  Its  formation  begins  to- 
ward the  close  of  the  growing  season. 

Structure  of  Epicotyl. 

The  epidermal  cells,  at  first  square  or  pentagonal  in  cross 
section,  become  at  length,  considerably  flattened.  There  are 
numerous  short,  curved  and  pointed  hairs.  A  poorly  developed 
hypoderma  is  present  in  year-old  material.  The  cells  of  the 
cortex  are  all  about  the  same  size. 

The  endodermis  was  distinguished  only  in  young  material. 
The  cells  are  small  and  contain  starch.  The  perimedullary 
zone,  cortex  and  phloem  have  at  a  later  time,  small  amounts  of 
starch. 


Ramaley :    HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.     119 


The  phloem,  in  the  youngest  material  examined,  forms  a 
complete  ring.  There  are  about  six  xylem  bundles.  These 
soon  fuse. 

Considerable  masses  of  stereom  develop  at  the  periphery  of 
the  phloem  forming  a  broken  sheath. 

The  pith  is  large-celled.  There  is  present  a  definite  peri- 
medullary  zone  of  small  cells  containing  starch.  The  cell  walls 
are  lignified. 

Cork  is  produced  in  the  outermost  hypodermal  layer  as  in 
other  species  of  Rhamnus  (cf.  Moeller  [1882],  pp.  292  et  seq.). 

Comparison  of  Structure  of  Hypocotyl  and  Epicotyl. 

The  hypocotyl  has  a  thicker  cortex  than  the  epicotyl ;  it  does 
not  have  a  hypoderma  ;  epidermal  hairs  are  absent ;  the  scleren- 
chymatous  ring  found  in  the  epicotyl  is  here  absent. 

The  endodermis  of  the  hypocotyl  remains  distinct  for  a 
greater  time  than  that  of  the  epicotyl.  The  former  region  has 
originally  four  vascular  bundles  ;  the  latter  has,  in  the  youngest 
material  examined,  a  zone  of  phloem 
and  about  six  xylem  bundles.  Cork 
formation  in  the  hypocotyl  is  endoder- 
mal  while  in  the  epicotyl  it  is  hypoder- 
mal. 

VITACE^E. 

Vitis  cordifolia. 


Rhammw 

purehians 


Structure  of  Hypocotyl. 

The  cells  of  the  epidermis  are  nearly 
square  in  cross  section,  but  become  at 
length  considerably  flattened.  Many 
of  them  are  somewhat  prolonged, 
forming  short,  blunt  papillae.  A  thick 
cuticle  is  present.  In  cross  section  it 
appears  minutely  notched. 

Three  or  four  of   the   outer    layers 
of  the  cortex  are  small-celled,  but  not  collenchymatous.     The 
cells  of  deeper  layers  are  larger  and  all  about  the  same  size. 

The  endodermis  is  small-celled,  and  is  for  a  long  time  read- 
ily distinguished  because  it  contains  starch.  Starch  is  gen- 
erally absent  from  the  other  tissues,  but,  at  a  later  time,  ap- 
pears in  the  pith. 


FIG.  17. 


120 


MINNESOTA    BOTANICAL    STUDIES. 


There  are  four  primary  vascular  bundles.  Other  secondary 
bundles  soon  become  intercalated  and  finally  complete  rings  of 
phloem  and  xylem  are  produced. 

A  single  group  of  sclerenchymatous  elements  is  formed  at 
the  outer  edge  of  each  primary  vascular  bundle.  By  the  end 
of  the  season  other  smaller  groups  are  also  present. 

The  pith  finally  becomes  very  small.  Sometimes,  by  the 
projection  into  it  of  two  of  the  vascular  bundles,  a  line  of  xylem 
extends  nearly  across  it.  There  is  no  perimedullary  zone. 

The  cork  cambium,  as  in  other  species  of  Vitis  (cf.  Flot 
[1889]),  is  formed  about  the  close  of  the  first  season  in  the 
pericycle. 

Structure  of  Epicotyl. 

The  epidermal  cells  are  square  in  cross  section,  becoming  at 
length  flattened.  The  cuticle  is  like  that  of  the  hypocotyl.  A 
well-differentiated  collenchymatous  hypoderma  is  present.  The 
other  cells  of  the  cortex  are  parenchymatous. 

The  endodermis  contains  starch  and  is,  therefore,  easily  dis- 
tinguished. Toward  the  end  of  the  season  starch  is  also  found 
in  the  pith.  The  number  of  primary  vascular  bundles  is  var- 
iable. Usually  there  are  more  than  eight.  At  an  early  time 
closed  zones  of  phloem  and  xylem  are  produced. 

A  broken  ring  of  sclerenchyma  is  developed  toward  the  end 
of  the  first  year  in  the  pericycle. 

The  pith  is  thin-walled  and  large- 
celled.  A  definite  perimedullary  zone 
was  not  distinguished. 

The  cork,  as  in  the  hypocotyl,  is 
of  pericyclic  origin  (  cf.  Moeller 
[1882],  p.  207). 

Comparison  of  Structure  of  Hypo- 
cotyl and  EpicotyL 
The  hypocotyl  has  a  thicker  cortex 
than  has  the  epicotyl.     It  has  but  four 
primary  vascular  bundles  instead  of 
eight   or    more.     It    has    four    large 
masses  of  sclerenchyma  in  the  peri- 
cycle and  a  few  smaller  ones  instead 
of  a  more  nearly  continuous  scleren- 
FIG.  18.  chymatous  ring. 


VitiS  cordifolia 


Ramaley :   HYPOCOTVL  AND  EPICOTVL  IN  WOODY  PLANTS.     121 

The  pith  of  the  hypocotyl  becomes,  at  length,  nearly  ob- 
literated. 

EL^EAGNACEjE. 

Elaeagnus  umbellata. 
Structure  of  Hypocotyl. 

The  cells  of  the  epidermis  are  more  or  less  oblong  in  cross 
section.  There  are  no  epidermal  hairs.  The  outermost  layer 
of  the  cortex  becomes  somewhat  thick-walled  but  not  collenchy- 
matous.  The  cortex  has  about  six  cell  rows. 

The  endodermis  is  small-celled.  It  remains  distinct  for  a 
time,  but  in  sections  of  material  gathered  at  the  end  of  the  season 
it  was  not  distinguished.  Very  little  starch  is  found  in  any  of 
the  tissues  save  in  the  endodermis. 

The  stele  is  originally  four-angled.  A  single  phloem  bundle 
and  two  xylem  bundles  are  placed  in  each  of  the  angles.  The 
phloem  soon  forms  a  closed  ring  while  the  xylem  bundles  fuse 
in  pairs  and  increase  in  size.  This  leaves  a  cruciform  pith. 
The  continued  growth  of  the  xylem  produces  a  complete  zone 
surrounding,  at  length,  a  circular  pith.  Secretion  cells  in  the 
phloem  are  numerous. 

At  the  outer  edge  of  the  xylem,  in  old  material,  are  a  few 
patches  of  sclerenchyma  forming  a  very  much  interrupted  ring. 

The  pith  is  of  considerable  extent  and  is  surrounded  by  a 
perimedullary  zone  of  small  cells  containing  starch. 

Cork  formation  takes  place  far  down  in  the  cortex. 

Structure  of  Epicotyl. 

The  epidermis  consists  of  cells  which  are  oblong  in  cross 
section  with  the  tangential  about  twice  the  radial  diameter  even 
in  very  young  material.  The  peculiar  stellate  trichome  struc- 
tures, well  known  in  this  genus,  are  abundant. 

The  outer  cortical  cells  are  nearly  circular  in  outline,  when 
seen  in  cross  section.  They  are  somewhat  smaller  than  the 
cells  of  the  epidermis.  The  inner  cortex  is  composed  of  large 
cells  which  eventually  are  very  much  flattened  owing  to  pres- 
sure of  the  growing  parts  within. 

The  endodermis  is  small-celled  and  contains  starch.  In  old 
material  it  was  not  recognized. 

The  stele  is  circular  from  the  first.  In  the  youngest  material 
examined  the  phloem  forms  a  complete  ring  surrounding  about 


122 


MINNESOTA    BOTANICAL    STUDIES. 


six  xylem  groups.  The  xylem  bundles  soon  fuse  producing  at 
the  end  of  the  season  quite  a  thick  zone. 

An  interrupted  sclerenchymatous  ring  is  developed  in  the 
pericycle. 

The  pith,  which  is  composed  of  large  thin-walled  elements, 
is  surrounded  by  a  narrow  small-celled  perimedullary  zone  con- 
taining starch. 

The  cork  is  formed  rather  late  in  the  season  in  the  outermost 
cortical  layer  as  in  other  species  of  Elaeagnus  (cf.  Moeller 
[1882],  p.  117). 

Comparison  of  Structure  of  Hypocotyl  and  Epicotyl. 

The  hypocotyl  is  without  the  trichome  structures  so  noticeable 
in  the  epicotyl ;  the  stele  is  at  first  four-angled  instead  of  cylin- 
drical ;  there  are  four  phloem  bundles  and  eight  xylem  bundles 
instead  of  a  ring  of  phloem  and  six  xylem  bundles. 

At  the  end  of  the  season  the  vas- 
cular tissue  is  alike  in  the  two  regions 
but  the  pericyclic  sclerenchyma  of  the 
hypocotyl  is  less  abundant.  Cork  is 
developed  in  the  inner  cortex  of  the 
hypocotyl  and  in  the  outermost  layer 
of  cortex  in  the  epicotyl. 

MYRTACE^). 

Eucalyptus  globulus. 

Structure  of  Hypocotyl. 

The  epidermal  cells,  at  first  oblong,, 
radially  elongated,  become  at  length  in 
cross    section     nearly    square.       The 
cuticle,  which  is  covered  with  eleva- 
tions,  appears,  when  young,  in  cross 
section  minutely  serrate. 
There  is  nohypoderma  differentiated,  but  the  outermost  layer 
of  the  cortex  is  smaller-celled  than  the  layers  below.     The  cor- 
tical cells  are  large.     They  become  flattened  toward  the  end  of 
the  season  by  the  growth  of  the  internal  tissues. 

The  endodermis  is  composed  of  small  cells  containing  starch  ; 
it  at  length  becomes  indistinguishable.  Starch  is  for  the  most 
part  absent  from  other  tissues.  Lysigenous  secretion  reservoirs- 
are  found  in  the  conjunctive  tissue. 


Elaeagnus 
umbellate 


FIG.  19. 


Ramaley :   IIYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS. 

The  stele  is  four-sided.  The  general  shape  of  the  hypocotyl 
sometimes  follows  that  of  the  stele  (cf.  Irmisch  [1876]).  There 
are  originally  four  narrow  curved  phloem  bundles  and  the  same 
number  of  small  xylem  bundles.  The  phloem  soon  forms  a 
closed  ring ;  the  xylem  bundles  increase  in  size,  leaving  for  a 
time  a  cruciform  pith ;  but  eventually  the  xylem  also  forms  a 
complete  ring  and  the  pith  is  cylindrical. 

Four  small  groups  of  sclerotic  cells  make  their  appearance 
in  the  pericycle  about  the  time  that  the  phloem  ring  is  first 
formed.  These  groups  eventually  become  somewhat  broken 
up  and  numerous  groups  of  fibers  appear  in  the  phloem  ar- 
ranged in  three  or  more  interrupted  circles. 

The  pith  is  large-celled.  It  is  small  in  amount  even  from  the 
first.  No  definite  perimedullary  zone  was  distinguished. 

Cork  formation,  according  to  Flot  [1890],  is  cortical  or  peri- 
cyclic. 

Structure  of  Epicotyl. 

The  cells  of  the  epidermis  are  at  first  more  nearly  square  in 
cross  section  than  those  of  the  hypocotyl.  They  at  length  be- 
come very  much  elongated  in  a  tangential  direction. 

The  cortex  is  large-celled ;  the  cells  of  the  outer  layer  are 
rather  small.  No  hypoderma  is  differentiated.  Numerous 
lysigenous  secretion  sacs  are  present. 

The  endodermis  is  thin-walled  ;  the  cells  are  small  and  con- 
tain starch. 

The  stele  is  at  first  somewhat  quadrangular  and  becomes  at 
length  elliptical,  in  cross  section.  In  the  youngest  material  ex- 
amined the  phloem  forms  a  closed  ring.  The  number  of  xylem 
bundles  is  somewhat  variable  ;  these  are  so  disposed  that  the  pith 
is  generally  at  first  somewhat  cruciform. 

The  pericycle  becomes,  at  length,  largely  sclerenchymatous  ; 
numerous  interrupted  rings  of  bast  fibers  begin  to  appear  but  are 
only  slightly  thickened  the  first  year. 

The  pith,  at  first  cruciform,  becomes  somewhat  quadrangular. 
There  is  a  perimedullary  zone  (fide  Flot  [1893]).  An  inter- 
nal cambium  produces  a  ring  of  phloem  just  outride  the  pith. 
This  is  mentioned  by  DeBary  [1884].  A  few  sclerotic  cells 
were  noted  at  the  inner  limit  of  the  internal  phloem. 

According  to  Flot  [1890]  cork  formation  is  sub-epidermal. 


124 


MINNESOTA    BOTANICAL    STUDIES. 


Comparison  of  Structure  of  Hypocotyl  and  Epicotyl. 

The  hypocotyl  is  more  nearly  cylindrical  than  the  epicotyl ; 
its  epidermis  less  flat,  its  stereom  is  better  developed  the  first 
year.  Stem  internodes  above  the  epicotyl  are  square. 

The  primary  structure  of  the  hypocotyl  is  like  that  of  Te- 
coma.  There  are  four  xylem  bundles  and  four  phloem  bundles. 
In  the  youngest  epicotyl  examined  the  phloem  forms  a  complete 
ring  and  there  is  a  variable  number  of  xylem  bundles. 

No  internal  phloem  was  recognized 
in  the  hypocotyl  although  it  is  quite  dis- 
tinct in  the  epicotyl.  The  cork  of  the 
former  region  is  pericyclic  or  cortical 
in  origin,  that  of  the  latter  subepi- 
dermal. 

BIGNONIACE^E. 


Eucalyptus 

"lobulus 


FIG.  20. 


Tecoma  radicans. 
Structure  of  Hypocotyl. 

The  epidermal  ceils  are  rectangular 
in  outline  when   seen  in  cross  section. 
From    being  originally   radially  elon- 
gated   they    are,     at    length,     nearly 
square.     The  cuticle  in  cross  section 
appears  minutely  serrate.     There  are 
a  few  short,  simple,  epidermal  hairs. 
The  cortex  is  of  loose  parenchyma,  generally  about  six  layers 
in  thickness.     There  is  no  differentiated  hypoderma. 

The  endodermal  cells  are  smaller  than  the  cells  of  the  cortex. 
Originally  they  are  irregularly  hexagonal  in  outline,  but  toward 
the  end  of  the  year  they  become  elliptical  and  have  slightly 
thickened  walls.  Starch  is  entirely  absent,  except  in  the  en- 
dodermal region  until  nearly  the  close  of  the  first  year,  when  it 
appears  especially  in  the  pith. 

The  stele  is  originally  very  small  and  somewhat  four-sided, 
containing  four  xylem  bundles  and  four  phloem  bundles.  The 
latter  are  next  the  pericycle.  They  alternate  with  the  xylem 
bundles.  These  are  presumably  the  four  "principal  bundles" 
found,  according  to  Hovelacque  [1888],  in  all  Bignoniaceous 
stems.  At  quite  an  early  stage  the  phloem  and  xylem  form 
closed  rings. 


Ramaley :   HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.     125 

About  the  time  that  this  is  apparent  four  small  groups  of 
sclerenchyma  make  their  appearance  in  the  pericycle  just  out- 
side the  original  xylem  bundles.  Each  of  these  is  composed  of 
only  six  to  ten  cells  with  extremely  thick  walls.  Later  numer- 
ous isolated,  lignified  sclerenchymatous  cells  appear  in  the  outer- 
most layer  of  the  cortex  ;  a  few  also  are  found  in  the  phloem 
area. 

The  pith  is  large-celled  and  thin-walled.  The  formation  of 
a  cambium  layer  in  the  small-celled  perimedullary  region  begins 
some  time  before  the  close  of  the  year.  This  will  be  further 
noticed  in  the  description  of  the  epicotyl. 

Cork  formation  takes  place  in  the  second  cortical  layer,  *'.  <?., 
in  the  cell  layer  immediately  below  the  sclerenchyma  which  is 
thus  eventually  lost. 

Structure  of  Epicotyl. 

The  epidermis  has  a  well-marked  cuticle  which,  in  cross  sec- 
tion, appears  minutely  notched.  The  cells  seen  in  cross  sec- 
tion are  about  square,  but  become  tangentially  elongated  toward 
the  end  of  the  season.  There  are  occasional  short  epidermal 
hairs. 

The  cells  of  the  outermost  layer  of  the  cortex  are  somewhat 
smaller  than  those  of  deeper  layers.  These  are  considerably 
flattened.  Although  at  first  of  about  even  thickness  through- 
out, the  cortex  soon  grows  in  thickness  at  four  equidistant 
points  giving  the  epicotyl  a  quadrangular  prismatic  shape. 

The  endodermis,  which  in  young  stages  is  distinct,  at  length 
becomes  indistinguishable  owing  to  displacement  and  crowding 
of  the  cells  caused  by  growth  in  the  lower  layers.  Starch, 
though  present  in  the  endodermal  region,  is  found  only  in  very 
small  amount  in  the  cortex  and  pith  until  the  close  of  the  sea- 
son. Even  then  the  cells  are  not  closely  packed  with  it. 

Even  in  the  youngest  stage  examined,  i.  e.,  second  stage  of 
our  arbitrary  division,  the  phloem  and  xylem  form  complete 
rings. 

The  outermost  cortical  layer  toward  the  end  of  the  season  be- 
comes largely  sclerenchymatous,  although  here  and  there  are 
cells  with  but  slightly  thickened  walls.  Certain  cells  of  the 
pericycle,  at  first  but  slightly  differentiated,  form,  at  length, 
groups  of  very  thick- walled  cells. 

The  pith  is  composed  of  large  cells  with  thin,  unlignified 


126 


MINNESOTA    BOTANICAL    STUDIES. 


walls.  It  is  found  to  be  practically  in  the  center  of  the  section, 
though  Pedicino  [1876]  found  that  when  the  plant  climbs  the  pith 
is  eccentric.  About  the  time  that  the  seedling  has  developed  one 
internode  above  the  epicotyl  a  medullary  cambium  has  begun 
to  form  in  the  epicotyl.  This  produces  xylem  without  and 
phloem  next  the  pith.  Considerable  masses  of  phloem  may 
thus  be  formed.  This  peculiar  cambium  in  Tecoma  was  noted 
by  Sanio  in  1864  and  fully  described  later  by  De  Bary  [1884]. 
Young  branches  of  the  plant  were  studied  by  these  investigators 
who  did  not  examine  seedlings. 

Cork  formation,  as  previously  described  by  Moeller  [1882] 
for  young  branches,  takes  place  in  the  second  layer  of  the  cor- 
tex. The  cork  cells  are  nearly  square  in  cross  section. 

Comparison  of  the  Structure  of  Hypocotyl  and  Epicotyl. 

Although  both  hypocotyl  and  epicotyl  are  originally  cylindri- 
cal, only  the  former  remains  so,  the  latter  developing  four 
thickened  areas  which  make  it  somewhat  quadrangular. 

The  four  small  groups  of  scleren- 
chyma  in  the  pericycle  of  the  hypo- 
cotyl are  represented  in  the  other  region 
by  a  considerable  number  of  smaller 
groups  forming  an  interrupted  ring. 
The  endodermis  remains  distinct 
in  the  hypocotyl  for  a  longer  time  than 

//      /^^\       \       in  the  epicotyl.     The  medullary  cam- 
bium is  formed  later  and  is  less  active. 


Tc'coma 

radicans 


Catalpa  speciosa. 

Structure  of  Hypocotyl. 

The  epidermis  consists  of  small  cells, 
square  in  cross  section.  Short,  blunt 
epidermal  hairs  are  rather  numerous. 
No  hypoderma  is  developed ;  all  the 
cortical  cells  are  thin-walled. 
The  endodermis  remains  distinct  for  a  long  time.  It  consists 
of  thin-walled  cells  which  are  but  slightly  smaller  than  the  cells 
of  the  cortex.  Starch  is  present,  from  the  first,  in  the  endoder- 
mis and  toward  the  close  of  the  season  appears  sparingly  dis- 
tributed in  medullary  rays  and  cortex ;  it  is  apparently  absent 
from  the  pith. 


FIG.  21. 


Ramalcy :   IIYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.     127 

The  stele  is  small ;  in  cross  section  it  is  circular.  There  are 
at  first  six,  seven  or  eight  conjoint  vascular  bundles  arranged  in 
a  circle.  Eight  is  probably  the  original  number,  but  fusions 
often  take  place  between  adjoining  bundles.  .Complete  zones  of 
xylem  and  phloem  are  formed  at  an  early  stage.  About  this 
time  four  small  groups  of  sclerenchyma  appear  in  the  pericycle  ; 
they  are  equidistant.  These  eventually  become  somewhat 
divided,  and  other  cells  of  the  pericycle  become  sclerotic,  so 
that  a  number  of  small  groups  of  stereom  are  found  in  this  area. 

The  pith  is  small  in  amount.  The  cells  are  large,  with  thin, 
slightly  lignified  walls.  There  is  a  perimedullary  zone  of  small 
cells  containing  starch. 

Cork  formation  takes  place  in  the  outermost  layer  of  cortical 
cells. 

Structure  of  Epicotyl. 

The  epidermal  cells  at  first  are  oblong  in  cross  section ; 
the  long  axis  is  at  right  angles  to  the  periphery  of  the  section. 
Later  the  shape  is  more  nearly  square.  There  are  many  straight, 
blunt  epidermal  hairs. 

The  first  two  or  three  cell  layers  of  the  cortex  are  collenchy- 
matous.  The  other  layers  are  rather  small-celled  parenchyma. 

The  endodermis,  though  at  first  distinct  on  account  of  the 
presence  of  starch  in  its  cells,  was  not  recognized  in  older  ma- 
terial. Starch  is  absent  from  the  other  tissues  in  the  early 
stages,  but  is  at  length  found  in  the  cortex,  phloem-,  medullary 
rays  and  perimedullary  zone. 

Toward  the  end  of  the  first  year  a  narrow,  much  interrupted 
ring  of  sclerenchyma  appears  at  the  outer  edge  of  the  phloem. 
The  cells  are  small  with  very  narrow  lumen. 

The  number  of  vascular  bundles  is  somewhat  variable. 
About  twenty  is  the  usual  number.  These  soon  unite  to  form 
zones  of  xylem  and  phloem. 

The  pith  is  large,  the  cells  thin-walled  parenchyma.  There 
is  a  perimedullary  zone  of  small  cells  containing  starch. 

Cork  arises  in  the  outermost  hypodermal  layer,  as  it  does  in 
the  stem  of  Catalpa  catalpa  (cf.  Moeller  [1882],  p.  184). 

Comparison  of  Structure  of  Hypocotyl  and  Epicotyl. 

The  epidermis  of  the  hypocotyl  has  fewer  and  shorter  hairs 
than  that  of  the  epicotyl.  The  former  region  has  no  hypo- 


128 


MINNESOTA    BOTANICAL    STUDIES. 


Catalpa 
speciosa 


derma,  though  in  the  epicotyl  a  distinct  collenchymatous  zone  is 

developed. 

The  endodermis  of  the  hypocotyl  remains  distinct  for  a  much 
longer  time  than  that  of  the  epicotyl ; 
the  stele  has  about  eight  vascular 
bundles,  instead  of  twenty  or  more ; 
sclerenchyma  is  first  disposed  in  four 
groups  instead  of  a  considerable  num- 
ber. 

The  pith  of  the  hypocotyl  is  much 
smaller  in  amount  than  that  of  the 
epicotyl. 

RUBIACE^. 

Cephalanthus  occidentalis. 
Structure  of  Hypocotyl. 

The  cells  of  the  epidermis  are  ob- 
long or  somewhat  hexagonal  in  cross 
section.     About    every    fifth    or   sixth 
FIG.  22.  cell  is  elongated  radially  and  pointed, 

projecting  somewhat  beyond  the  gen- 
eral line  of  cells.  These  might  be  described  as  extremely 
short  hairs.  This  characteristic  feature  continues  for  a  consid- 
erable length  of  time. 

No  distinct  hypoderma  is  formed.  The  cells  of  the  two  or 
three  outer  layers  of  the  cortex  are  rather  thick-walled  but  not 
collenchymatous.  The  cortex  is  loose  with  numerous  large  in- 
tercellular spaces. 

The  endodermis  is  large-celled.  It  remains  distinct  through 
the  first  year  although  the  cells  become  at  length  very  much 
flattened.  They  contain  starch.  Starch  is  later  found  in  the 
various  parenchymatous  tissues. 

The  stele  is  circular  in  cross  section.  There  are  originally 
four  phloem  bundles  and  an  equal  number  of  xylem  bundles  ; 
they  are  grouped  in  pairs.  The  xylem  and  phloem  soon  form 
closed  zones,  the  xylem  encroaching  upon  the  pith  which,  at 
the  close  of  the  first  season  is  almost  entirely  obliterated.  The 
hypocotyl  thus  assumes  a  root-like  structure — "  rhizelle "'  of 
Van  Tieghem  [1891]. 

The  cork  is  of  epidermal  origin. 


Ramaley :   HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.     129 


Structure  of  Epicotyl. 

The  epidermal  cells  are  rectangular  in  cross  section.  The 
radial  is  the  long  diameter  at  first  but  eventually  the  two  diame- 
ters are  nearly  equal.  Many  of  the  cells  are  prolonged  to 
form  pointed  hairs  which  are  about  three  times  as  long  as  the 
ordinary  cells  of  the  epidermis. 

A  narrow  collenchymatous  hypoderma  is  developed ;  this 
shades  off  gradually  into  the  ordinary  cortex,  which  is  quite 
extensive. 

The  endodermis  is  rather  large-celled,  the  cells  resembling 
those  of  the  cortex  but  containing  starch.  The  endodermis  re- 
mains distinct  throughout  the  first  year.  Starch,  which  is  at 
first  absent  from  the  other  tissues,  becomes,  at  length,  dis- 
tributed through  all  the  parenchymatous  elements. 

The  stele,  originally  elliptical  in  cross  section,  follows  the 
general  shape  of  the  epicotyl.  Eventually  the  epicotyl  be- 
comes cylindrical  as  does  also  the  stele.  In  the  youngest  ma- 
terial examined,  the  phloem  forms  a  complete  zone  surrounding 
a  ring  of  about  six  xylem  bundles,  which  soon  fuse  to  form  a 
closed  ring. 

A  few  of  the  cells  of  the  pericycle  become  sclerotic  after  a 
time.  These  are  generally  isolated  ;  not  aggregated  in  groups. 

The  pith  becomes  quite  small ;  it  is  surrounded  by  a  well- 
developed  small-celled  perimedullary  zone  whose  elements  con- 
tain starch. 

The  cork,  like  that  of  the   hypo- 
cotyl,  arises  in  the  epidermis. 


Comparison  of  Structure  of  Hypo- 
cotyl  and  Epicotyl. 

The  hypocotyl  differs  from  the  epi- 
cotyl in  its  shorter  epidermal  hairs,  in 
the  absence  of  a  true  hypoderma  and 
in  the  much  looser  parenchyma  of  its 
cortex. 

The  structure  of  the  stele  is  also 
very  different,  the  hypocotyl  having 
originally  four  phloem  bundles  and 
four  xylem  bundles,  while  in  the  epi- 
cotyl the  phloem,  even  in  the  youngest 


Cephalanthus 
occidentals 


FIG.  23. 


130  MINNESOTA    BOTANICAL    STUDIES. 

stage  examined,  forms  a  closed   ring   surrounding  six    xylem 
bundles. 

At  the  end  of  the  first  year  the  structure  of  the  two  regions  is 
essentially  the  same  except  that  the  hypocotyl  is  without  pith  or 
differentiated  hypoderma. 


GENERAL  CONCLUSIONS. 

The  following  summary  and  conclusions  are  based  on  the 
facts  shown  in  the  foregoing  pages.  It  is  not  intended  to  re- 
peat here  all  the  points  which  are  there  given,  but  merely  to 
bring  together  under  appropriate  headings  the  most  important 
facts  of  structure  of  the  hypocotyl  and  epicotyl  in  the  plants 
studied. 

General  Shape  of  Hypocotyl  and  Epicotyl. — In  cross  sec- 
tion the  hypocotyl  is  usually  circular  in  outline,  the  epicotyl  is, 
however,  not  infrequently  hexagonal  in  outline  and  somewhat 
flattened.  The  hypocotyl  has  usually  in  early  stages  much  the 
greater  diameter. 

Comparison  of  the  Epidermis  of  Hypocotyl  and  Epicotyl. — 
The  epidermal  cells  of  both  regions  when  seen  in  cross  section 
appear  at  first  square  or  radially  elongated.  After  a  time, 
however,  they  became  elongated  tangentially,  being  stretched 
by  the  growth  of  the  stelar  tissues  and  not  continuing  to  divide. 
In  quite  young  stages  some  plants  have  in  the  epicotyledonary 
region,  epidermal  cells  which,  in  cross  section,  appear  tangen- 
tially elongated,  viz.  :  Ulmus  americana,  Liriodendron  tulip- 
ifera,  Butneria  florida,  Cercis  canadensis.  Trichome  struc- 
tures are  usually  fewer  and  less  complex  in  the  hypocotyledon- 
ary  region,  e.  g".,  Ulmus  americana,  Broussonetia  papyrifera, 
Butneria  florida,  Robinia  pscudacacia,  Ailanthus  glandulosa, 
Catalpa  speciosa,  Cephalanthus  occidentalis.  In  the  following 
species  the  epicotyl  has  trichome  structures  but  they  are  absent 
from  the  hypocotyl :  Celtis  occidentalis,  Toxylon  pomiferum, 
Gleditsia  triacanthos,  Ptelea  trifoliata,  Rhamnus  purshiana, 
El&agnus  umbellata,  Tecoma  radicans. 

Hypoderma  tn  the  Hypocotyl  and  Epicotyl. — Only  one  of  the 
species  examined  has  a  definite  collenchymatous  hypoderma  in 
both  regions.  This  is  Butneria  Jlorida.  The  following  plants 
have  a  hypoderma  in  the  epicotyl,  but  not  in  the  hypocotyl :  Celtis 


Ramaley :   HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.     131 

cccidentalis^  Broussonetia  papyrifera,  Liriodendron  tulipifera, 
Menispermum  canadcnse,  Gleditsia  triacanthos,  Robinia  pseu- 
dacacia,  Ptelea  trifoliata,  Ailanthus  glandnlosa,  Rhamnus 
purs/iiana,  Catalpa  spcciosa,  Cephalanthus  occidentals. 

Comparison  of  Cortex  of  Hypocotyl  and  Epicotyl. — The 
cortical  cells  of  the  hypocotyl  are  nearly  always  much  larger 
than  those  of  the  epicotyl.  This  is  so  commonly  the  case 
that  various  species  need  not  here  be  specially  mentioned ;  a 
good  example  is  Parkinsonia  aculeata.  The  cortex  of  the  hy- 
pocotyl is  thicker  than  that  of  the  epicotyl. 

Comparison  of  Endodermis  and  Pericycle  in  Hypocotyl  and 
Epicotyl. — The  endodermis  in  the  hypocotyl  is,  as  a  rule,  more 
distinct  and  persists  longer  than  that  of  the  epicotyl.  Its  cells 
contain  starch.  Pericycle  is  well  developed  in  the  hypocotyl, 
usually  consisting  of  two  or  more  layers  of  small  cells. 

Typical  Structure  of  the  Stele  of  the  Hypocotyl.  The  stele 
is  usually  somewhat  quadrangular.  As  a  rule  there  are  four 
phloem  bundles  and  four  xylem  bundles.  The  phloem  and 
xylem  may  be  in  contact  or  they  may  be  separated  by  a  small 
amount  of  un differentiated  parenchyma.  In  the  latter  case 
each  phloem  area  is  either  directly  outside  of  a  xylem  area 
(when  the  phloem  may  be  spoken  of  as  opposite  the  xylem),  or 
the  phloem  bundles  are  removed  from  the  xylem  by  greater  or 
less  angular  distances  (alternate  arrangement).  Using  the 
foregoing  terminology  the  arrangement  of  bundles  may  be  de- 
scribed as  opposite  in  the  following  :  Liriodendron  tulipifcra, 
Menispermum  canadense,  Butneria  jlorida,  Cercts  canadensis, 
Amorpha  fruticosa,  Ptelea  trifoliata,  Ailanthus  glandulosa, 
Schinus  molle,  Berchemia  racemosa,  Rhamnus  purshiana,  Vitis 
cordifolia,  Cephalanthus  occidentalis.  It  is  alternate  in  Eu- 
calyptus globulus  and  Tecoma  radicans. 

Certain  modifications  of  the  more  usual  type  just  described 
would  best  be  noted  separately.  There  are  four  xylem  bundles 
and  eight  phloem  bundles  in  Robinia  psetidacacia.  In  Parkin- 
sonia aculeata,  Gleditsia  triacanthos  and  Elaagnus  umbellata 
there  are  eight  xylem  bundles  and  four  phloem  bundles.  The 
xylem  bundles  soon  fuse  together  in  pairs  in  the  last  two  named 
species.  In  Celtis  occidentalis  and  Toxylon  pomiferum,  the 
phloem  forms  two  crescent-shaped  areas  while  the  arrangement 
of  the  xylem  is  normal. 

Unusual  Structure  of  the  Stele  of  the  Hypocotyl. — In  certain 


132  MINNESOTA    BOTANICAL    STUDIES. 

species  the  stele  of  the  hypocotyl  does  not  have  the  typical 
structure  just  mentioned,  the  number  and  arrangement  of  vascu- 
lar  bundles  being  different.  Ulmus  americana  has  two  xylen: 
crescents  and  numerous  small  phloem  bundles ;  Broussonetic 
•papyrifera  has  a  root-like  structure  ;  in  Catalpa  speciosa  then 
are  about  eight  vascular  bundles. 

Primary  Structure  of  the  Stele  of  the  Epicotyl. — In  shape  the 
stele  of  the  epicotyl  is  often  originally  somewhat  hexagonal 
though,  as  in  the  case  of  the  hypocotyl,  becoming  at  length 
cylindrical.  As  is  well  known  there  are  usually  from  six  tc 
very  many  vascular  bundles.  Sometimes  the  phloem  is  com- 
pletely fused  into  a  closed  zone  even  in  very  young  stages. 

Arrangement  of  Sclerenchyma  in  the  Hypocotyl.  The  scler- 
enchyma of  the  hypocotyl  first  appears  as  four  masses  in  the 
peri  cycle  in  Toxlyon  pomiferum,  Liriodendron  tulipifera,  Cer- 
cis  canadensis,  Gleditsia  triacanthos,  Robinia  pseudacacia,  Ail 
anthus  glandulosa,  Vitis  cordifolia,  Eucalyptus  globulus,  7  e- 
coma  radicans  and  Catalpa  speciosa.  In  the  plants  just  namec 
this  original  disposition  of  the  sclerenchyma  becomes  alterec 
either  by  the  intercalation  of  parenchymatous  elements  in  the 
areas  of  sclerenchyma  or  by  the  development  of  sclerenchyrm 
at  other  points.  In  the  following  plants,  however,  there  is 
practically  no  change  in  the  sclerenchyma  during  the  first  yeai 
and  the  four  original  masses  remain  to  the  end  of  the  season 
Parkinsonia  aculeata,  Amorpha  fruticosa,  Ptelea  trifoliata, 
Berchemia  racemosa. 

Comparison  of  the  Hypocotyl  and  Epicotyl  -with  Refer  enct 
to  the  Distribution  of  Sclerenchyma.  Commonly  the  scleren- 
chyma in  the  two  regions  becomes,  at  the  close  of  the  first  grow- 
ing season,  equally  well  developed  and  similarly  arranged.  Ex- 
ceptions to  this  rule  will  now  be  noted.  Sclerenchyma  is  absen 
from  the  hypocotyl  of  Menispermum  canadense,  Butneria  flo- 
rida  and  Rhamnus  purshiana  although  present  in  the  epicotyl 
In  Ptelea  trifoliata  and  Berchemia  racemosa  at  the  close  of  the 
first  year  there  is  a  greater  development  of  sclerenchyma  in  the 
epicotyl  than  in  the  hypocotyl.  The  reverse  of  this  conditior 
obtains  in  Celtis  occidentalis  and  Eticalyptus  globulus.  Onl} 
scattered  sclerenchyma  in  small  amount  was  recognized  ir 
either  region  in  year-old  material  of  Broussonetia  papyrifera^ 
in  Cephalanthus  occidentalis  a  few  only  of  the  pericyclic  cells 
of  the  epicotyl  become  sclerotic. 


Ramalcy :    HYPOCOTYL  AND  EPICOTYL  IN  WOODY  PLANTS.     133 

Cork  Formation  in  Hypocotyl  and  Epicotyl.  The  cork  cam- 
bium is  developed  in  the  layer  of  cells  next  below  the  epi- 
dermis in  both  hypocotyl  and  epicotyl  of  Celtis  occidentalis^ 
Liriodcndron  tulipifera,  Butncria  florida,  Ptelca  trifoliata^ 
Ailanthtis  g'landulosa,  Catalpa  spcciosa.  In  Cephalanthus  oc- 
cidentalis  it  is  of  epidermal  origin  in  both  regions.  In  the 
following  species  cork  formation  is  sub-epidermal  in  the  epi- 
cotyl but  the  cork  is  produced  in  deeper  cell  layers  of  the  hypo- 
cotyl :  Ulnms  americana,  Toxylon  -pomiferum,  Broussonetia 
papyrifera,  Rhanmus  purshiana,  El&agnus  umbellata*  Euca- 
lyptus globulus.  In  Cercis  canadensis,  Gleditsia  triacanthos^ 
Amorphafruticosa,  Robinia  pseudacacia,  Vitis  cordifolia  and 
Tecoma  radicans  cork  formation  in  the  epicotyl  is  cortical,  while 
in  the  hypocotyl  it  takes  place  in  some  cases  in  the  same  cell 
layer,  in  other  cases  in  deeper  layers.  Details  are  given  in  the 
previous  descriptions  for  the  separate  species. 

Pith  and  Perimedullary  Zone  of  Hypocotyl  and  Epicotyl. 
The  pith  of  the  hypocotyl  is  smaller  than  that  of  the  epicotyl, 
sometimes  it  becomes  nearly  obliterated,  e.  g.,  Cephalanthus  oc- 
cidcntalis.  The  perimedullary  zone  is  sometimes  not  distin- 
guished in  the  hypocotyl  though  present  in  the  epicotyl,  e.  g., 
Parkinsonia  aculeata,  Gleditsia  triacanthos,  Eucalyptus  globu- 
lus. The  opposite  condition  is  found  in  Cercts  canadensis  and 
Berchemia  racemosa.  More  often  where  a  perimedullary  zone 
is  recognized  it  is  equally  developed  in  both  hypocotyl  and  epi- 
cotyl. 

Structure  of  Hypocotyl  and  Epicotyl  at  the  close  of  the  first 
years  growth. — Owing  to  secondary  changes  the  two  regions, 
though  at  first  quite  dissimilar  in  structure,  may  come  to  be  very- 
much  alike.  The  xylem  and  phloem  always  form  closed  rings  ; 
the  endodermis  often  becomes  indistinguishable ;  the  cells  of 
the  cortex  become  flattened.  The  differences  of  pith,  perime- 
dullary zone  and  sclerenchyma  have  already  been  given. 

Condensed  Summary. 

Although  secondary  changes  may  cause  a  great  resemblance 
in  the  structure  of  hypocotyl  and  epicotyl,  the  two  regions  are, 
in  their  primary  structure,  essentially  dissimilar. 

The  epidermis  of  the  hypocotyl  is  more  often  without  trichome 
structures,  the  cortex  is  thicker  and  composed  of  larger  cells, 


134  MINNESOTA    BOTANICAL    STUDIES. 

the  endodermis  is  more  distinct  and  persists  for  a  greater  length 
of  time,  the  pith  is  smaller,  sclerenchyma  is  often  less  well-de- 
veloped and  differently  arranged  and  a  hypoderma,  so  common 
in  the  epicotyl,  is  nearly  always  absent.  Cork  formation  in 
the  hypocotyl  begins  either  in  the  same  cell-layer  that  it  does 
in  the  epicotyl  or  in  a  deeper  layer,  never  in  a  more  superficial 
one. 

Starch  is,  as  a  rule,  distributed  in  the  same  way  in  both 
regions.  It  is  usually  present  in  the  endodermis  in  the  early 
stages  but  does  not  appear  in  other  tissues  until  the  plant  has 
developed  foliage  leaves,  in  considerable  number. 

As  to  the  structure  of  the  stele  it  may  be  said  that  in  the 
hypocotyl  there  are  usually  four  primary  vascular  bundles. 
The  exact  disposition  of  the  phloem  and  xylem  elements  is 
subject  to  some  variation.  Occasionally  there  are  more  than 
four  bundles.  In  the  epicotyl  the  vascular  bundles  are  from 
six  to  eight  or  very  many. 

The  hypocotyl  does  not  have  a  root-like  structure. 

EXPLANATION  OF  PLATES. 

Plate  V.  Drawings  of  cross  sections  to  show  the  primary  stelar 
structure  of  the  hypocotyl  of  Ulmus  americana,  Celtis  occidentalism 
1 ' oxylon  pomiferum  and  Broussonetia  papyrifera. 

Plate  VI.  Drawings  of  cross  sections  to  show  the  primary  stelar 
structure  of  the  hypocotyl  of  Menispermum  canadense,  Butneria 
florida,  Amorpha  fruticosa,  Robinia  pseudacacia  and  Ptelea  tri- 
foliata. 

Plate  VII.  Drawings  of  cross  sections  to  show  the  primary  stelar 
structure  of  the  hypocotyl  of  Ailanthus  glandulosa,  Schinus  molle, 
Berchemia  racemosa,  Eucalyptus  globulus,  Catalpa  speciosa  and 
Cephalanthus  occidentalis. 

Plate  VIII.  Photographs  of  cross  sections  to  show  the  primary 
structure  of  the  hypocotyl.  I.  Ulmus  americana,  2.  Liriodendron 
tulipifera,  3.  Parkinsonia  aculeata,  4.  Cercis  canadensis,  5- 
Berchemia  racemosa,  6.  Vitis  cordifolia,  *].  Eucalyptus  globtdus, 
8.  Tecoma  radicans. 

The  drawings  were  all  outlined  with  the  aid  of  a  camera  lucida. 
The  magnification  used  was  about  five  hundred  diameters.  For  pub- 
lication the  drawings  have  been  reduced  to  one-half  their  original  size. 
The  magnification  used  in  making  the  photographs  was  from  fifty  to 
eighty  diameters.  They  have  been  slightly  reduced.  All  drawings 
and  photographs  were  made  by  the  author  from  his  own  preparations. 


Ramaley :    HVPOCOTYL  AND  EPKOTYL  i\  WOODY  PLANTS.     1.S5 


BIBLIOGRAPHY   OF   ARTICLES    TO  WHICH    REFERENCE    is  MADE  IN 
THE  FOREGOING  PAPER. 

Dangeard,  P.  A.  Le  mode  d'union  de  la  tige  et  de  la  racine  chez 
les  angiospermes.  Comptes  rendus.  2e  Setn.  107:  287.  1888. 

Dangeard,  P.  A.  Rech.  sur  la  mode  d'union  de  la  tige  et  de  la 
racine  chez  les  dicotylcdones.  Le  Botaniste.  i :  75.  1889. 

De  Bary,  A.  Compar.  Anat.  of  the  Vegetative  Organs  of  the 
Phanerogams  and  Ferns.  English  translation.  1884. 

Flot,  Leon.  Sur  la  region  tigellaire  des  arbres.  Comptes  rendus. 
ler  Sem.  108  :  306.  1889. 

Flot,  Leon.  Rech.  sur  la  structure  comparee  de  la  tige  des  arbres. 
Rev.  gen.  de  Bot.  a:  17-32,  66-77,  122-136.  1890. 

Flot,  LCon.     Rech.    sur   la   zone  pe'rimedullaire  de  la  tige.     Ann. 
Sc.  Nat.  Bot.  VII.  18:  37.      1893. 

Gerard,  R.  Rech.  sur  la  structure  de  1'axe  au-dessous  des  feuilles 
seminales  chez  les  dicotylcdones.  Comptes  rendus  go  :  1295.  1880. 

Gerard,  R.  Rech.  sur  le  passage  de  la  racine  a  la  tige.  Ann.  Sc. 
Nat.  Bot.  VI.  ii :  279.  iSSi. 

Goldsmith,  Sophie.  Beitriige  zur  Entwicklungsgeschichte  der  Fi- 
brovasalmassen  im  Stengel  und  in  der  Hauptwurzel  der  Dicotyledonen. 
Inaug.  Diss.  Zurich,  pp.  48.  1876. 

Herail.  Rech.  sur  1'  anat.  comp.  de  la  tige  des  dicotyl£dones. 
Ann.  Sc.  Nat.  Bot.  VII.  2:  203.  1885. 

Hovelacque,  Maurice.  Characteres  anatomique  genereaux  de  la 
tige  des  Bignoniacees.  Bull.  Soc.  d'Etudes  scientif.  de  Paris,  nme 
annee.  1888. 

Irmisch,  Th.  Einige  Beobachtungen  an  Eucalyptus  globulus  Lab. 
Zeitschr.  f.  ges.  Naturwiss.  48: — 1876.  (Just.  Bot.  Jahresbericht  4 : 
440.  1878.) 

Klebs.  Beitrage  zur  Morph.  und  Biol.  der  Keimung.  Pfeffer's 
Untersuchungen  aus  dem  Bot.  Inst.  zu  Tubingen.  1 :  536.  1885. 

Moeller,  J.     Anatomic  der  Baumrinden.      1882. 

Pedicino,  N.  A.  Studii  sulla  struttura  e  sulla  maniera  di  accres- 
cersi  di  alcuni  fusti  di  piante  dicotiledoni.  Annuar.  della  R.  Scuola 
Super.  d'Agricoltum  di  Portici.  1876.  (Just.  Bot.  Jahresbericht, 
Abt.  L,  6:  41.  1880.) 

Sanio,  C.  Notiz  iiber  Verdickung  des  Holzkorpers  auf  der  Mark- 
seite  bei  Tecoma  radicans.  Bot.  Zeit.  22:  61.  1864. 

Trecul.  Des  vaisseaux  propres  dans  les  Terebinthinees.  Comptes 
rendus  65  :  17.  1867. 

Troschel,  J.  Untersuchungen  iiber  das  Mestom  im  Holze  der 
dicotylen  Laubbiiume.  Verhandl.  des  Bot.  Ver.  der  Provinz  Bran- 
denburg, 21 :  78.  1879. 


136  MINNESOTA    BOTANICAL    STUDIES. 

Van  Tieghem,  Ph.  Sur  les  canaux  secreteurs  des  Liquidambarees 
et  des  Simarubacees.  Bull.  Soc.  Bot.  France  II.  6:  247.  1884. 

Van  Tieghem,  Ph.  Sur  la  limite  de  la  tige  et  de  la  racine  dans 
1'hypocotyle  des  Phanerogames.  Morot  Journ.  de  Bot.  5  :  425.  1891. 

Williams,  J.  Lloyd.  The  sieve  tubes  of  Calycanthus  occidentalis, 
Ann.  of  Bot.  8:  367.  1894. 

Woronin,  M.  Ueber  d.  Ban  d.  Stammes  von  Calycanthus.  Bot. 
Zeit.  18:  177.  1860. 


OL.   II. 


MINNESOTA 


Ulnius  americana 


Toxylon  pomiferum 


L    STUDIES. 


PART    II. 


Celtis  occidentalis 


Hroussonetia  papyrifera 


VOL.    II. 


MINNES 


Menispermum  canadense 


Butneria  florida 


IICAL    STUDIES. 


Ptelea  trifoliate 


OL.   II. 


MINNESOTA 


Ailauthus  glaudulosa 


Schinus  molle 


ICAL    STUDIES. 


Eucalyptus  globulus 


Ja  speciosa 


Cephalanthus  occidentalis 


VOL.   II. 


MINNE 


Ulmus  araericana 
Berchemia  racemosa 


Parkinsonia  aculeata 
Cercio  canadensis 


NICAL    STUDIES. 


Eucalyptus  globulus 
Tecoma  radicans 


Liriodendron  tulipifera 
Vitis  eordilblia 


X.  CONTRIBUTION  TO  THE  LIFE-HISTORY  OF 

RUMEX. 


BRUCE  FINK. 


Introduction. — The  preliminary  work  leading  to  the  present 
paper  was  nearly  all  done  in  the  botanical  laboratories  of  the 
University  of  Minnesota  during  the  summers  of  1896  and  1897. 
This  work  included  a  study  of  the  macrosporangia  and  their  con- 
tents in  Bursa  bursa--pastoris  (L.)  Britton,  Silene  antirrhina 
L.,  Polygonum  erectum  L.,  Rumex  acetosella  L.,  Rumex  salici- 
folius  Weinm.,  and  Rumex  verticillatus  L. 

At  the  close  of  the  season  of  1897,  I  had  about  an  equal  num- 
ber of  good  preparations  of  the  last  two  species  and  had  learned 
that  Rumex  verticillatus  is  a  much  more  favorable  plant  for 
study  than  Rumex  salicifolius  because  of  the  larger  size  of  the 
structures  to  be  investigated.  Consequently,  during  the  latter 
part  of  August,  1897,  after  previous  study  of  the  former  plant 
had  enabled  me  to  select  methods  which  seemed  best  adapted  to 
my  purpose,  a  large  number  of  flowers  and  buds,  selected  to 
represent  various  stages  of  development  of  the  gametophyte, 
were  preserved  for  future  study.  During  the  summer  of  1898 
over  three  thousand  macrosporangia  were  sectioned,  and  the 
drawings  presented  with  this  paper  were  reproduced  from  the 
preparations  that  gave  the  best  results.  Rumex  salicifolius 
was  also  further  studied  in  1898,  and  the  series  of  slides  of  this 
plant  is  nearly  as  complete  as  that  for  Rumex  verticillatus.  As 
the  two  plants  gave  very  similar  results,  except  for  size  of 
structures,  I  have  not  thought  it  necessary  to  multiply  figures  by 
giving  a  full  series  for  both  plants.  The  figures  then  are,  in 
the  main,  drawn  from  preparations  from  the  latter  plant ;  those 
from  the  former  being  introduced  only  when  the  equivalent 
phases  of  development  were  not  found  in  the  other  plant,  or 
were  poorly  exhibited  in  the  preparations. 

The  plants  of  the  genus  studied  are  not  well  adapted  to  the 


138  MINNESOTA    BOTANICAL    STUDIES. 

study  of  nuclear  mechanism  and  phenomena  because  of  the 
small  size  of  the  cells  and  contained  nuclei,  and  I  have  conse- 
quently confined  myself  to  other  phases  of  the  subject.  Nor 
have  I  found  anything  in  the  study  that  would  give  additional 
evidence  as  to  the  nature  of  various  structures  within  the  em- 
bryo sac  so  that  my  work  resolves  itself  into  a  description  of 
the  gametophyte  and  some  comparative  studies. 

Under  the  subject  stated  I  shall,  for  the  sake  of  relationships, 
begin  with  the  archesporium,  which  is  the  last  term  in  the 
sporophytic  generation  and  also  include  stages  immediately  fol- 
lowing the  establishment  of  the  sporophyte. 

So  far  as  I  know  this  is  the  first  work  done  on  the  female 
gametophyte  of  ffumex,  or  of  any  plant  within  the  Polygona- 
ceae,  except  Polygonum  divaricatum,  which  has  been  investi- 
gated by  Strasburger. 

I  am  under  obligations  to  Professor  Conway  MacMillan  for 
helpful  suggestions  as  to  technique  and  interpretation  of  struc- 
ture and  for  access  to  the  literature  of  the  subject. 

Origin  of  the  Macrospore. — At  about  the  usual  stage  in  the 
development  of  the  macrosporangium,  an  axial  hypodermal 
cell  at  the  summit  of  the  nucellus  begins  to  enlarge  and  soon 
contains  a  larger  nucleus  and  denser  cytoplasm  than  the  sur- 
rounding cells  ( Fig.  i ).  This  cell  constitutes  the  archespor- 
ium, and  in  all  instances  examined,  only  one  cell  showed  this 
archesporial  nature.  The  archesporium  in  plants  may  develop 
directly  into  the  macrospore ;  it  may  itself  become  a  sporogen- 
ous  cell  ( mother  cell )  and  divide  into  a  number  of  potential 
macrospores ;  or  more  commonly  it  divides  first  into  a  tapetum 
and  a  sporogenous  cell,  each  of  which  may  divide,  forming 
a  cellular  tissue.  In  JRumex  the  last  order  of  development  is 
followed.  After  increasing  considerably  in  size  (comp.  Figs,  i 
and  2),  the  archesporium  divides  by  a  periclinal  wall  into  the 
inner  sporogenous  cell  (the  mother  cell  of  the  macrospore),  and 
the  outer  hypodermal  tapetum  (Fig.  2).  This  apparently  pro- 
tective tapetum  proceeds  next  to  divide,  sometimes  by  a  peri- 
clinal wall  (Fig.  4),  but  no  doubt  much  more  commonly  by 
an  anticlinal  (Fig.  3).  In  no  instance  did  I  observe  more  than 
four  tapetal  cells  derived  from  the  primary  tapetum  and  often 
only  three,  one  of  the  two  derived  from  the  primary  tapetum 
apparently  failing  to  divide.  Indeed,  sometimes  only  one  tape- 
tal cell  could  be  distinguished  at  a  period  of  development  which 


Fink:     CONTRIBUTION    TO    THE    LIFE-HISTORY    OF    RUMKX.      139 

led  to  the  suspicion  that  the  tapetum  may  sometimes  fail  to  di- 
vide. Fig.  5  shows  four  tapetal  cells  and  the  sporogenous  cell 
below  somewhat  more  elongated  than  usual  with  its  nucleus  un- 
usually near  its  upper  end.  That  this  elongated  cell  is  the 
mother  cell  of  the  macrospore  instead  of  the  macrospore  itself 
is  proved  by  the  outline  of  the  inner  seed  coat,  which  shows  its 
early  development  as  compared  with  its  more  advanced  condi- 
tion before  the  macrospore  is  produced  (Fig.  10).  Fig.  6  rep- 
resents a  single  tapetum  lying  above  a  dividing  mother  cell,  and 
this  may  be  the  original  tapetum  which  has  failed  to  divide, 
though  a  cell  lying  almost  directly  below  it,  and  hence  not 
shown  in  the  figure,  may  have  been  a  second  tapetal.  Fig.  3 
shows  a  typical  mother  cell  apparently  about  ready  to  divide, 
and  showing  two  tapetals,  neither  of  which  would  be  likely  to 
divide  again. 

After  the  formation  of  the  protective  cap  of  tapetal  cells  and 
the  enlargement  of  the  mother  cell  of  the  macrospore,  the  latter 
divides  as  usual,  among  the  Archechlamydeae  at  least,  into  a  row 
of  four  potential  macrospores.  It  is  here  that  our  subject  proper 
really  opens  since  this  mother  cell,  in  which  the  reduction 
of  chromosomes  takes  place,  stands  between  the  sporophytic 
and  gametophytic  generations,  connected  morphologically  with 
the  former  and  physiologically  with  the  latter.  As  the  mother 
cell  divides  the  nucleus  lies  longitudinally  at  or  a  little  above 
the  center  of  the  cell  (Fig.  6).  As  elsewhere  in  these  studies, 
the  number  of  chromosomes  could  not  be  made  out,  but,  in  all 
probability  through  failure  to  get  a  complete  view  of  individual 
chromosomes,  there  really  seemed  to  be  twenty-four  in  this  nu- 
cleus. The  two  cells  resulting  from  this  division  were  not  seen, 
but  from  the  position  of  the  dividing  nucleus  in  several  instances 
observed,  it  may  be  assumed  that  the  division  is  into  two  cells 
of  approximately  equal  size  as  observed  by  Strasburger*  in  Po- 
lygonum  divaricatum.  This  dividing  nucleus  of  the  mother 
cell  of  the  macrospore  was  about  as  large  as  that  of  the  macro- 
spore  itself  (Fig.  9)  and,  being  like  the  latter  a  nucleus  of  a 
large  well-fed  and  consequently  somewhat  inactive  cell,  was  ap- 
parently a  long  time  in  dividing.  The  reduction  of  chromosomes 
is  supposed  to  be  a  process  involving  more  time  than  is  com- 
monly occupied  in  mitotic  division  of  nuclei ;  and  this  process 
doubtless  also  added  further  to  the  time  occupied  by  this  mother 

*  Strasburger,  E.  Die  Angiospermen  und  die  Gymnospermen,  p.  5,  1879. 


140  MINNESOTA    BOTANICAL    STUDIES. 

cell  nucleus  in  dividing,  so  that  it  was  seen  dividing  much  more 
frequently  than  the  equally  large  and  apparently  as  well-fed 
nucleus  of  the  macrospore. 

As  stated  above  the  two  cells  resulting  from  the  division  of 
the  macrospore  mother  cell  were  not  seen  in  the  resting  condi- 
tion, though  the  four  potential  macrospores  resulting  from  their 
division  were  frequently  found.  The  two  cells  were  seen  once 
with  their  nuclei  dividing,  but  the  nuclei  were  indistinct  because 
of  improper  staining.  The  two-celled  condition,  which  was 
not  seen,  is  doubtless  a  very  transient  phase,  the  upper  of  the 
two  cells  almost  immediately  cutting  off  a  small  cell  from  its 
lower  end,  and  the  lower  of  the  two  likewise  dividing  at  once 
into  two  cells,  a  large  one  below  and  a  small  one  above  ( Fig.  7  ). 
This  four-celled  stage  seems  to  be  constant  and  of  quite  long 
duration  as  it  was  frequently  seen  as  represented  in  Fig.  7  or  as 
in  Fig.  10.  In  the  latter  the  lower  more  successful  cell  which 
is  to  become  the  fertile  macrospore  has  begun  to  absorb  the 
other  three  potential  macrospores  for  its  own  nourishment. 
This  may  be  seen  by  observing  the  decrease  in  amount  of  cyto- 
plasm contained  in  each  of  the  three  cells  as  compared  with  the 
same  three  in  Fig.  7  and  the  swelling  of  the  softened  periclinal 
cell  walls  between  any  two  of  them  and  between  the  lowest  one 
and  the  absorbing  cell  below.  The  next  observed  develop- 
mental condition  was  that  in  which  the  three  cells  were  all  ab- 
sorbed except  possibly  a  refractive  cytoplasmic  cap  at  the  sum- 
mit of  the  absorbing  cell,  which  more  probably  represents  a 
nearly  absorbed  tapetal  cell  (Fig.  8).  This  brings  us  to  the 
macrospore  as  shown  in  the  figure. 

Germination  of  the  Macrospore. — When  first  formed  the 
macrospore  has  very  nearly  the  shape  of  the  four  cells  replaced 
and  shows  the  nucleus  at  the  center  surrounded  by  cytoplasm, 
while  the  upper  and  lower  ends  are  each  occupied  by  a  large 
vacuole.  Though  I  cannot  account  for  this  apparent  poverty 
in  cytoplasm  at  this  time,  the  condition  seems  to  be  typical.  As 
the  macrospore  increases  in  size  by  the  absorption  of  tapetal 
cells  and  those  cells  of  the  macrosporangium  which  surround  its 
upper  lateral  wall,  cytoplasm  increases  in  amount  (Fig.  9).  In 
the  figure  the  nucleus  is  lying  in  its  usual  longitudinal  direction 
about  the  middle  of  the  macrospore  while  dividing.  Before  the 
nucleus  divides,  the  macrospore  increases  considerably  in  size 
by  the  absorption  and  pressing  upon  surrounding  tissues  (comp. 


Fink :   CONTRIBUTION  TO  THE  LIFE-HISTORY  OF  RUMEX.    141 

Figs.  8  and  9).  The  cavity  of  the  macrospore,  which  I  shall 
now  designate  by  the  usual  name  of  embryo  sac,  continues 
to  increase  in  size  as  it  approaches  the  condition  shown  in  Fig. 
ii  by  the  continual  absorption  and  pressure  upon  surrounding 
cells.  The  relative  position  of  cytoplasm  and  vacuoles  in  Fig.  9 
is  hardly  normal,  the  nucleus  of  the  macrospore  more  com- 
monly lying  along  the  central  longitudinal  axis  of  the  spore. 
In  another  preparation  showing  the  dividing  nucleus  of  the  mac- 
rospore, the  nucleus  was  in  this  more  usual  position,  and  no 
vacuole  was  seen.  Fig.  11  shows  the  two  nuclei  derived  from 
the  nucleus  of  the  macrospore  and  two  tapetals  and  two  other 
sporangial  cells  nearly  absorbed.  Of  these  two  nuclei  it  is 
quite  common  to  find  the  lower  one  larger,  probably  as  a  result 
of  better  nourishment,  and,  apparently  consequently  giving  rise 
in  division  to  a  larger  number  of  chromosomes. 

The  condition  as  to  chromosomes  could  not  be  studied,  but  I 
noticed  relative  sizes  carefully.  As  instances  of  difference  in 
size  of  the  two  nuclei,  the  researches  of  Sargant,*  Mottierf  and 
Guignardij:  may  be  cited.  I  examined  a  number  of  the  embryo 
sacs  showing  the  two  nuclei,  and  it  would  seem  that  the  lower 
nucleus  becomes  very  slightly  larger  than  the  upper  (Figs,  n 
and  12) .  The  slight  difference  may  not  be  constant,  and  indeed 
in  the  closely  related  Polygonum  divaricatum  Strasburger§  shows 
the  upper  nucleus  larger  than  the  lower.  This  stage  of  develop- 
ment was  quite  frequently  observed,  but  the  next,  in  which  two 
nuclei  appear  in  each  end  of  the  sac,  was  only  seen  twice  (Figs. 
13  and  14) .  The  first  of  the  two  figures  shows  the  probable  posi- 
tion of  the  two  nuclei  during  division  as  a  persistent  spindle  was 
seen  between  the  anterior  pair.  Then  the  anterior  pair  seems 
to  result  from  the  division  of  a  nucleus  lying  transversely  in  the 
anterior  end  of  the  sac  and  the  posterior  pair  from  a  posterior 
nucleus  lying  longitudinally  in  the  sac.  This  position  of  the 
two  nuclei  while  dividing  is  the  common  one  in  plants  so  far  as 
I  can  ascertain.  Fig.  14  represents  a  later  stage,  in  which  the 


*Sargant,  Ethel.  The  Formation  of  the  Sexual  Nuclei  in  Lilium  martagon. 
I.  Oogenesis.  Annals  of  Bot.  10  :  464.  S.  1896. 

tMottier,  David  M.  Ueber  das  Verhalten  der  Kerne  bei  der  Entwicklung  des 
Embryosack  und  die  Vorgange  bei  der  Befruchtung.  Jahrbucher  fur  wissen- 
schaftliche  Botanik  31  :  132.  1897. 

JGuignard,  L.  Nouvelles  Etudes  sur  le  Fecundation.  Ann.  des  Sci.  naturelles 
Botanique  VII.  14:  187.  1891. 

§Strasburger,  E.     1.  c.  Fig.  15. 


142  MINNESOTA    BOTANICAL    STUDIES. 

nuclei  have  changed  position  somewhat.  A  difference  in  size 
has  frequently  been  observed  here  also,  in  some  other  plants  the 
lower  pair  being  larger.  I  may  cite  Guignard*  again  and  also 
Mottier.f  However,  the  last  named  investigator  only  states  that 
the  nuclei  forming  the  lower  pair  are  much  larger  just  before 
dividing,  distinctly  stating}:  that  at  an  earlier  period  the  four  are 
all  of  equal  size.  I  did  not  observe  this  phase  of  development 
often  enough  to  be  certain  that  the  size  is  the  same  at  all  times 
in  Rtimex;  but  it  seems  from  the  instances  observed  that,  though 
coming  from  the  nuclei  differing  somewhat  in  size,  the  four  are 
so  nearly  of  the  same  size  at  all  times  that  any  difference  would 
be  difficult  to  detect.  During  the  change  to  the  four-nucleate  con- 
dition the  sac  increases  somewhat  in  size,  as  may  be  seen  by 
comparing  Figs.  13  arid  14  with  Fig.  12. 

The  development  from  the  four-nucleate  to  the  eight-nucleate 
condition  must  be  even  more  rapid  than  that  from  the  two-nucle- 
ate to  the  four-nucleate  phase,  for  I  was  neither  able  to  find  the 
two  tetrads  in  position,  nor  the  division  of  the  four  nuclei  leading 
to  its  establishment.  The  nearest  approach  to  it  was  observed 
in  Rumex  salicifolius  (Fig.  15),  when  the  polar  nuclei  were  ap- 
proaching. The  condition  represented  in  Fig.  15  is  an  espe- 
cially interesting  step  in  the  life-history  of  the  gametophyte  of 
Rumex  because  of  departure  from  the  usual  conditions  and  espe- 
cially from  the  nearly  related  Polygonum.  On  examining  the 
figures  of  Guignard,  Vesque,  Strasburger,  Ward  and  others,  I 
find  that  in  fully  five-sixths  of  their  drawings  they  show  cell 
walls  about  the  three  anterior  cells  before  the  polars  have  fused. 
Strasburger  figures  for  Polygonum  divaricatum  §  three  nuclei 
enclosed  in  cellular  membranes  and  one  free  nucleus  in  each 
tetrad  even  before  the  two  polars  begin  to  approach  each  other. 
My  own  observations  on  Polygonum  erectum  L.  (Fig.  16), 
Bursa  bursa-pastoris  (  L.)  Britton  and  Silcne  antirrhina  L.  gave 
the  same  results  so  far  as  the  anterior  end  of  the  sac  is  con- 
cerned, though  the  evanescent  antipodals  of  Bursa  were  not 
satisfactorily  studied  in  this  respect.  Methods  which  brought 
out  these  walls  in  the  three  plants  named  above  should  show 
them,  if  present  in  Rumex,  yet  in  this  genus  I  find  all  the  an- 

*Guignard,  L.,  1.  c.  188. 
tMottier,  David  M. ,  1.  c.  137. 
JMottier,  David  M.,  1.  c.  136. 
§  Strasburger,  E.     1.  c.  Fig.  17. 


Fink  :     CONTRIBUTION    TO    THE    LIFE-HISTORY    OF    RUMEX.      143 

terior  three  cells  free  while  the  polars  are  approaching  each  other 
in  Rumex  salicifolius  ( Fig.  15).  When  the  polars  have  met, 
or  are  fusing  in  this  plant,  the  cellulose  walls  seem  to  begin  to 
form,  and  a  suggestion  of  such  a  structure  may  occasionally  be 
seen  (Figs.  17  and  20)  about  the  nuclei  of  the  synergidae.  In 
Rumex  verticillatus,  even  shortly  after  the  definitive  nucleus  is 
formed  (Fig.  18),  I  could  only  distinguish  a  suggestion  of  a 
beginning  of  formation  of  a  membrane  about  the  lower  end  of 
the  egg  while  the  sister  nuclei  which  form  the  synergidae  were 
yet  free.  In  Rumex  salicifolius  the  synergidae  form  earlier 
than  in  Rumex  vcrticillatus  and  are  more  regular  in  form  in  the 
former  plant  (compare  Fig.  24  with  Figs.  19  and  23).  So  far 
as  investigation  has  proceeded  then,  cell  walls  appear  about  the 
three  anterior  nuclei  (excluding  the  anterior  polar)  before  the 
meeting  of  the  polar  gametes  in  Polygonum;  while  in  Rumex 
the  walls  appear  after  the  meeting  of  the  gametes,  or  even  after 
their  fusion  to  form  the  zygote  constituting  the  definitive  nucleus. 
The  matter  is  one  of  some  taxonomic  interest,  which  can  only 
be  solved  by  a  laborious  study  of  all  the  species  of  the  two  genera 
with  reference  to  this  particular  problem  in  developmental  history. 
Regarding  conditions  presented  in  Figs.  15,  17  and  20  some 
further  statements  are  necessary.  In  these  stages  I  was  able  to 
note  no  difference  in  size  of  the  three  anterior  nuclei,  while  at 
later  periods  the  nucleus  of  the  egg  had  increased  in  size  so 
that  it  was  larger  than  those  of  the  synergidas  (Figs.  18  and  19). 
So  far  as  I  could  ascertain,  the  nuclei  of  the  three  antipodals 
were  smaller  than  the  three  just  discussed  at  all  periods.  Fig. 
17  represents  the  three  antipodals  as  best  seen  in  Rumex  salici- 
folius^ and  here  their  smaller  size  can  be  seen  distinctly.  The 
antipodals  are  difficult  of  observation  at  all  times  because  of 
their  lying  in  the  small  caacum-like  prolongation  of  the  embryo 
sac  so  that,  except  in  very  thin  sections,  other  cellular  struc- 
tures of  the  surrounding  macrosporangium  almost  always  par- 
tially or  completely  obscure  them.  In  the  sac  of  the  same  plant 
showing  a  slightly  earlier  stage  of  development  (Fig.  15),  only 
the  deeply-stained  nucleoli  could  be  seen  through  the  overlying 
tissues  of  the  macrosporangium,  neither  the  nuclear  membrane 
nor  the  cellular  wall  about  each  of  the  three  nuclei  appearing. 
In  this  plant  the  three-celled  antipodal  area  was  found  persisting 
in  the  latest  phases  studied  after  the  establishment  of  the  em- 
bryonic sporophyte  (Fig.  33)  and  seems  to  be  quite  constant, 


144  MINNESOTA    BOTANICAL    STUDIES. 

though  in  the  sac  represented  in  Fig.  24  I  suspect  that  there 
were  really  no  walls  about  the  antipodal  nuclei.  However, 
here  again  overlying  tissues  may  have  obscured  them.  Rumex 
verticillatus  gave  less  satisfaction  in  the  study  of  the  antipodal 
region.  In  only  one  instance  were  three  antipodals  seen  con- 
stituting a  three-celled  mass  of  tissue  (Fig.  21).  This  figure 
represents  the  posterior  end  of  a  mature  sac.  In  other  instances 
various  conditions  of  cell-wall  formation  were  shown  in  this  re- 
gion before  the  establishment  of  the  sporophyte  (Figs.  23,  27 
and  28).  Sometimes  more  than  three  nuclei  could  be  distin- 
guished within  this  antipodal  area  (Figs.  23  and  28).  After 
fertilization  these  cells  seem  to  disintegrate  more  or  less  in  this 
plant,  and  no  such  typical  structure  was  found  persisting  as  is 
figured  for  Rumex  salicifolius  (Fig.  33).  The  degenerating 
condition  of  the  antipodal  region  just  after  fecundation  is  shown 
in  Fig.  27,  which  is  the  lower  end  of  the  sac  presented  in  Fig. 
26.  Here  I  was  only  able  to  make  out  a  highly  refractive  area 
with  neither  walls  nor  nuclei.  As  to  the  time  when  the  cellular 
membranes  appear  around  the  antipodal  nuclei,  I  am  not  able 
to  state  certainly  because  of  the  difficulty  encountered  in  inves- 
tigation, but  it  seems,  from  the  facts  presented  and  other  ob- 
served phenomena,  that  in  Rumex  salicifolius  they  form  earlier 
than  those  about  the  three  nuclei  in  the  anterior  end  of  the  sac. 
In  Rumex  verticillatus  walls  were  not  detected  till  the  time  of 
maturity  of  the  embryo  sac  when  the  typical  three  celled  condi- 
tion (Fig.  21)  presented  itself,  or  that  of  three  or  more  nuclei 
within  a  common  wall  (Fig.  23).  The  antipodal  area  in  Rumex 
salicifolius  seems  to  agree  substantially  with  the  third  of  the 
four  types  proposed  by  Coulter*  of  "  three  comparatively  per- 
manent cells  not  notable  for  size  or  activity  and  usually  associ- 
ated with  a  sac  decidedly  narrowed  at  the  antipodal  end."  The 
antipodals  of  Rumex  verticillatus  are  not  so  permanent,  but 
doubtless  should  be  classed  here  also  as  should  those  of  Poly- 
gonum  erectum. 

The  size  of  the  two  polars  in  plants  may  differ  as  well  as 
their  place  of  fusion.  Schaffner  finds  the  upper  one  larger  in 
Alisma  -plantago\  and  Sagittaria  variabilis.\  Also  he  finds  in 

*Coulter,  John  M.  Contribution  to  the  Life-History  of  Ranunculus.  Bot. 
Gaz.  25:  80-8 1.  F.  1898. 

t  Schaffner,  John  H.  The  Embryo  Sac  of  Alisma  plantago.  Bot.  Gaz.  21  : 
126.  Mr.  1896. 

+  Schaffner,  John  H.  Contribution  to  the  Life  History  of  Sagittaria  varia- 
bilis.  Bot.  Gaz.  23  :  255.  Ap.  1897. 


Fink  :     CONTRIBUTION    TO    THE    LIFE-HISTORY    OF    RUMEX.     145 

the  last  plants  that  the  two  polars  fuse  in  the  lower  part  of  the 
sac,  the  larger  gamete  strangely  enough  traveling  further  than 
the  smaller  and  thus  showing  greater  activity.  Mottier  *  finds 
them  of  about  the  same  size  in  Senccio  aureus  and  that  they 
fuse  in  the  anterior  end  of  the  sac  just  below  the  egg,  the  lower 
of  the  two  equal-sized  polars  thus  showing  greater  activity.  In 
Rumex,  and  all  of  the  Polygonaceas  so  far  as  studied,  the  two 
polar  nuclei  are  of  about  the  same  size  (Figs.  15,  17  and  20) 
and  fused  at  or  near  the  center  of  the  sac. 

During  the  passage  from  the  four-nucleate  to  the  eight-nucle- 
ate condition,  the  sac  increases  considerably  in  size  (comp.  Figs. 
13  and  15),  and  the  increase  in  size  is  even  more  noticeable  as 
the  sac  matures  (comp.  Figs.  15  and  24). 

The  Mature  Embryo  Sac. — The  mature  sac  contains  typically 
the  usual  seven  nuclei,  though  in  Rumex  verticillatus^  as  stated 
elsewhere,  the  number  in  the  antipodal  region  may  vary,  pro- 
ducing a  corresponding  variation  in  the  total  number  contained 
in  the  sac.  In  Rumex  verticillatus  the  synergidas  are  some- 
what irregular  in  form  (Figs.  19  and  23),  while  in  Rumex 
salicifolius  they  are  more  regular  in  outline  (Fig.  24).  The 
egg  usually  lies  in  contact  with  the  lower  part  of  one  synergid 
in  both  plants  (Figs.  19,  23  and  24),  and  its  nucleus  is  much 
larger  than  those  of  the  synergidae.  All  three  nuclei  are  en- 
closed in  definite  walls  at  this  time.  The  definitive  nucleus  is 
much  the  largest  nucleus  in  the  sac  (Figs.  18,  23,  24,  25  and 
28).  In  Rumex  verticillatus  it  usually  approaches  the  egg 
after  formation  and  lies  close  to  it  till  the  time  for  fecundation 
approaches  (Fig.  23)  when  it  commonly  recedes  somewhat 
(Figs.  25  and  26).  In  Rumex  salicifolius  it  usually  lies  at 
some  distance  from  the  egg  in  the  mature  sac  (Fig.  24)  though 
it  was  once  seen  nearer  in  an  earlier  stage  (Fig.  22).  In  Ru- 
mex salicifolius  it  may  be  assumed  that  the  antipodals  are  in  the 
typical  three-celled  condition  at  maturity  as  they  were  observed 
in  this  condition  both  before  and  after  as  already  stated  (Figs. 
17  and  33).  As  before  noted  the  condition  of  the  antipodals  in 
Fig.  24  is  doubtful.  The  condition  of  the  antipodals  in  Rumex 
verticillatus  at  this  time  has  also  been  explained  above. 

In  shape  the  mature  sac  in  both  plants  differs  quite  widely 
from  that  of  Polygonum  divaricatum  as  shown  in  Strasburger's 

*  Mottier,  David  M.     On  the  Embryo  Sac  of  Senccio  aureus.     Bot.  Gaz.  18  : 
Fig.  I  and  p.  248.     Jl.  1893. 


146  MINNESOTA    BOTANICAL    STUDIES. 

figure,*  the  largest  part  being  near  the  anterior  end  and  the 
posterior  narrowed  end  being  quite  long  (Figs.  17,  23,  24,  etc.) 
while  his  figures  show  the  narrowed  posterior  caecum  dilating 
quite  abruptly,  making  the  posterior  half  of  the  mature  sac  quite 
as  large  as  the  anterior  half.  Polygonum  erect um  the  mature 
sac  is  much  more  like  JRumex  than  Polygonum  divaricatum  as 
figured  by  Strasburger. 

Fecundation,  and  the  origin  of  the  Sporophyte.  During  the 
period  of  fecundation  and  the  establishment  of  the  sporophyte, 
the  embryo  sac  continues  to  increase  in  size  rapidly,  as  may  be 
seen  by  comparing  Figs.  24  and  33  and  also  23  and  30,  making 
allowance  for  the  greater  reduction  of  Figs.  30  and  33. 

The  first  evidence  of  a  pollen  tube  approaching  or  already 
present  in  the  sac  is  the  disappearance  of  one  of  the  syner- 
gidse  (Fig.  25).  When  actually  present  the  tube  is  an  easy 
object  to  detect  because  it  stains  more  deeply  than  surrounding 
tissues,  hence  in  those  instances  in  which  one  synergid  was 
breaking  down  while  no  tube  could  be  distinguished,  I  have 
concluded  that  the  tube  was  just  approaching  the  sac.  In  Fig. 
25 -the  tube  has  not  yet  discharged,  as  it  shows  two  nuclei  and 
the  end  is  intact,  and  it  lies  as  usual  beside  the  persistent  syn- 
ergid. The  definitive  nucleus  here  occupies  a  position  at  some 
distance  from  the  egg  as  it  frequently  does  at  this  time.  Here 
this  nucleus  is  in  a  resting  condition,  and  I  find  it  so  constantly 
up  to  this  time.  After  the  pollen  tube  has  discharged  the  re- 
maining synergid  disappears,  and  the  definitive  nucleus  soon 
divides  (Fig.  26).  Though  in  some  plants  the  definitive  nu- 
cleus seems  to  divide  sometimes  before  the  entrance  of  the  pol- 
len tube  into  the  sac,  its  presence  in  the  style  probably  furnish- 
ing the  necessary  stimulus,  it  does  not  divide  till  after  the 
entrance  of  the  tube  in  Rumex  verticillatus  and  probably  usually 
not  till  after  the  fecundation  of  the  egg.  In  the  discharge  of 
the  sperm  nucleus  the  tip  of  the  tube  is  ruptured  as  shown  in 
Figs.  26,  29,  30  and  35.  After  the  pollen  tube  has  discharged 
a  deeply  stained  spot  may  always  be  seen  within  the  tube  as 
shown  in  the  same  figures.  This  is  the  shrunken  second  and 
undischarged  sperm  nucleus.  The  fusion  of  the  sexual  nuclei 
was  only  seen  once,  and  in  that  instance  the  egg  was  badly 
distorted.  The  tube  in  this  case  was  closely  applied  to  the  egg 
though  the  figures  given  herewith  seem  to  indicate  that  this 

*  Strasburger,  E.     1.  c.  Fig.  21. 


J'"illk  :     CONTRIBUTION    TO    THE    LIFE-HISTORY    OF    RUMEX.      147 

is  not  the  usual  relation  during  fusion  of  these  nuclei.  The 
conditions  existing  in  Fig.  28  are  somewhat  of  an  enigma.  The 
sac  is  surely  larger  than  it  could  be  previous  to  fecundation,  but 
the  two  smaller  nuclei  in  the  anterior  end  look  like  those  of  the 
synergidae.  Also  the  definitive  nucleus  lies  near  the  egg  as  I 
uniformly  find  it  in  Rumex  verticillatus  before  fecundation.  I 
thought  for  a  time  that  I  had  here  a  pollen  tube  showing  one 
sperm  nucleus  within  and  the  other  discharged  and  entering 
the  egg.  However,  taking  into  account  the  appearance  of  the 
two  smaller  nuclei  and  the  restful  condition  of  the  egg,  I  have 
concluded  that  the  sac  is  one  whose  egg  failed  to  be  fertilized 
and  in  which  one  synergid  has  disintegrated  leaving  its  nucleus 
while  the  other  is  rapidly  dwindling.  If  this  is  true,  the  sac 
has  gone  on  increasing  in  size  the  same  as  if  the  egg  had  been 
fecundated.  The  next  stage  observed  after  that  already  ex- 
plained (Fig.  26)  is  that  represented  in  Fig.  30,  in  which  the 
tube  had  discharged,  the  first  two  endosperm  nuclei  had  divided 
and  the  spindles  were  persisting,  and  the  egg  was  evidently 
preparing  to  divide.  A  little  later  phase  was  also  found  (Fig. 
29),  in  which  the  tube  was  persisting  and  showed  the  broken 
end  beautifully,  the  egg  was  dividing  and  three  of  a  probable  four 
endosperm  nuclei  were  visible.  The  tube  frequently  persists  in 
both  plants  till  the  sporophyte  is  well  established  (Fig.  35),  and 
in  one  instance  an  undischarged  tube  was  seen  in  the  sac  of 
Rumex  verticillatus  after  the  embryo  was  well  established  and 
four  endosperm  nuclei  were  dividing  (Fig.  34).  This  tube, 
containing  two  sperm  nuclei,  is  a  second  one  which  entered  the 
sac  after  fecundation  had  been  accomplished. 

An  exhaustive  study  of  the  embryo  will  not  be  attempted ;  but 
I  shall  state  a  few  observed  facts  concerning  its  origin  and  early 
development,  making  no  use  of  the  terms  suspensor  and  pro- 
embryo,  but  designating  the  structure  from  the  beginning  as  the 
embryo.  The  first  division  then  of  the  embryo  occurs  soon 
after  the  egg  has  elongated  and  secreted  a  cellulose  wall  about 
its  base  to  attach  it  to  the  anterior  end  of  the  sac  and  is  trans- 
verse (Fig.  29).  The  second  and  third  divisions  were  also 
transverse  in  several  instances  observed  (Fig.  35),  and  the 
fourth  was  a  longitudinal  division  of  the  distal  cell  of  the  em- 
bryo (Fig.  31).  In  an  instance  observed  the  third  division  was 
transverse  and  closely  followed  by  a  longitudinal  dividing  the 
second  cell  from  the  distal  end  (Fig.  32).  During  its  early  de- 


148  MINNESOTA    BOTANICAL    STUDIES. 

velopment  the  embryo  is  not  always  attached  centrally  at  the 
anterior  end  of  the  sac,  nor  does  it  often  lie  in  an  exactly  longi- 
tudinal direction  in  the  sac. 

It  has  already  been  stated  that  the  definite  nucleus  in  Rumex 
does  not  divide  till  after  fecundation  of  the  egg,  an  observation 
based  upon  examination  of  two  or  three  hundred  sacs.  It  is 
worthy  of  note  that  the  endosperm  nuclei  observed  while  divid- 
ing in  any  given  sac  were  all  in  the  same  mitotic  phase  (Figs. 
30  and  34).  The  last  figure  shows  only  one  of  four  nuclei  ob- 
served dividing.  The  endosperm  nuclei  were  not  so  numerous, 
at  the  stages  studied,  in  Rumex  verticillatus  as  in  Rumex  salici- 
folius  (Fig.  33) ;  nor  were  they  yet  enclosed  in  cell  walls  in 
either  plant. 

Relation  of  the  Gametophyte  to  the  Macro  sporangium. — I 
have  studied  carefully  the  position  of  the  base  of  the  sporogen- 
ous  tissues  and  derived  embryo  sac  with  reference  to  lines  con- 
necting the  points  of  origin  of  the  seed  coats  in  order  to  as- 
certain how  much  of  the  enlargement  of  these  structures  is 
associated  with  a  downward  growth  and  consequent  crowding 
of  the  tissues  of  the  macrosporangium  and  how  much  is  accom- 
plished by  upward  growth,  keeping  pace  with  the  growth  of 
the  nucellus.  The  position  of  the  base  of  sporogenous  tissue, 
and  later  of  the  sac,  with  reference  to  these  lines  is  not  always 
quite  the  same  at  any  particular  stage  of  development ;  but  by 
comparative  study,  safe  conclusions  have  been  secured.  Be- 
tween the  base  of  the  archesporium  and  the  lines  connecting 
the  supposed  points  of  origin  of  the  future  seed  coat  are  five  or 
six  cells.  After  the  tapetum  is  cut  off  (Fig.  2),  there  are  only 
three  or  four  cells  between  the  base  of  the  mother  cell  of  the 
macrospore  and  the  lines.  By  the  time  of  division  of  the  tape- 
tum (Fig.  3)  the  base  of  the  mother  cell  is  within  two  or  three 
cells  of  these  lines.  During  this  time  the  nucellus  has  increased 
in  length  very  little,  its  increase  in  size  being  principally  in 
width.  Consequently,  this  fact,  together  with  the  relative  posi- 
tion of  the  base  of  sporogenous  tissue  and  the  lines  at  various 
stages  of  development,  indicates  that  the  sporogenous  tissue  has 
grown  downward  in  the  nucellus.  As  no  evidence  of  absorp- 
tion of  cells  was  seen  at  this  time,  I  conclude  that  this  down- 
ward growth  is  accomplished  by  crowding  downward  and  out- 
ward the  subjacent  layers  of  cells  of  the  macrosporangium. 
However,  the  effects  of  the  crowding  were  so  distributed  among 


CONTRIBUTION    TO    THE    LIFE-HISTORY    OP    RUMEX.      149 

several  layers  of  cells  that  they  were  scarcely  visible  in  any 
particular  cell.  After  the  mother  cell  has  reached  its  full  length, 
there  is  no  further  downward  growth,  the  further  increase  in 
length  of  the  sporogenous  tissues  and  subsequently  of  the  sac 
being  accompanied  by  a  proportionate  elongation  of  the  nucel- 
lus.  By  the  time  of  the  establishment  of  the  macrospore,  a 
thickening  of  the  walls  of  cells  in  the  chalazal  region  for  the 
support  of  structures  above  has  begun  in  a  layer  of  cells  ex- 
tending transversely  between  the  points  of  origin  of  the  inner 
seed  coat.  As  the  superimposed  structures  become  heavier,  the 
thickening  extends  to  several  layers  of  cells  below  the  ones  first 
thickened  and  gives  rise  to  quite  a  mass  of  thick-walled  tissue 
extending  entirely  across  the  chalazal  region. 

As  the  macrospore  matures  and  prepares  to  divide  (Fig.  9), 
absorption  of  tapetals  above  and  pressure  on  surrounding  cells 
of  the  upper  nucellus  becomes  evident  and  is  apparent  in  all 
subsequent  stages  of  development  of  the  gametophyte.  How- 
ever, the  swelling  of  the  cell  walls  of  the  upper  nucellar  tissue 
just  beneath  the  epidermis  as  observed  by  Strasburger*  in  Poly- 
gonum  divaricatum  as  a  result  of  absorption  I  have  not  seen 
either  in  Rumex  or  Polygonum. 

By  the  time  development  has  proceeded  to  the  condition  rep- 
resented in  Fig.  n,  the  subepidermal  cells  of  the  upper  end  of 
the  nucellus  have  all  been  absorbed,  and  from  this  time  on  till 
the  establishment  of  the  conditions  shown  in  Fig.  28,  or  possibly 
not  later  than  those  shown  in  Figs.  25  or  26,  the  increase  in 
size  is  due,  at  least  principally,  to  the  absorption  of  cells  of  the 
nucellus  surrounding  the  middle  portion  of  the  sac,  which  still 
continues  to  increase  in  size.  During  this  time  the  sac  is  in- 
creasing in  length,  and  since  there  has  been  no  further  sinking 
of  its  posterior  end  into  subjacent  tissues,  as  is  shown  by  the 
fact  that  its  lower  end  is  still  removed  from  the  lines  connecting 
the  points  of  origin  of  the  inner  seed  coat  by  two  or  three  cells, 
as  was  the  lower  end  of  the  mother  cell,  this  increase  in  length 
is  accompanied  by  an  equal  upward  growth  of  the  nucellus. 
Dividing  nuclei  were  seen  in  the  basal  region  of  the  nucellus,  both 
in  the  epidermis  and  in  the  sub-epidermal  cells,  up  to  the  latest 
stages  studied,  indicating  that  this  basal  portion  of  the  nucellus 
is  its  chief  region  of  growth  at  these  stages.  After  the  growing 
gametophyte  has  absorbed  all  the  sub-epidermal  tissues  of  the 

•Strasburger,  E.  1.  c.  Figs.  10  et  seq. 


150  MINNESOTA    BOTANICAL    STUDIES. 

upper  nucellus,  the  sac  does  not  cease  to  expand  laterally,  but 
presses  the  remaining  epidermis  of  this  portion  of  the  nucellus 
outward  as  it  still  further  increases  in  size.  These  epidermal 
cells  contain  cytoplasm  and  may  divide  even  after  the  cells  of 
the  inner  nucellus,  or  their  cytoplasmic  contents,  at  least,  have 
been  absorbed ;  but  the  increase  in  epidermal  surface  accom- 
panying the  continued  increase  in  size  of  the  sac  is  doubtless 
due  principally  to  increase  in  length  of  these  upper  epidermal 
cells  and  the  division  of  those  near  the  base  of  the  nucellus. 

Fig.  28  shows  certain  of  these  conditions  of  the  nucellus 
brought  about  by  absorption  of  its  tissues  by  the  growing  game- 
tophyte  and  by  its  own  growth.  All  of  the  epidermal  cells  ex- 
cept those  at  the  summit  show  elongation  in  the  direction  of 
upward  growth  of  the  nucellus.  All  of  the  epidermal  cells  ex- 
cept those  at  the  summit  are  also  well  filled  with  cytoplasm, 
indicating  activity.  The  lowest  sub-epidermal  cells  of  the 
nucellus  shown  in  the  figure  are  also  well  filled  as  were  the  cells 
of  five  or  six  layers  lying  between  trie  lowest  shown  in  the  fig- 
ure and  the  area  of  thickened  cells  in  the  chalazal  region.  In 
passing  upward  from  the  base  to  the  summit  of  the  nucellus,  we 
find  greater  and  greater  absorption  of  the  cytoplasmic  cell-con- 
tents and  finally  a  partial  breaking  down  of  the  cell  walls  and  a 
beginning  of  the  consequent  collapsing  of  the  cells.  In  the 
lower  portion  of  the  nucellus  the  absorption  by  the  gametophyte 
has  only  affected  the  layers  of  cells  near  the  sac,  while  in  the 
upper  portions  all  of  the  sub-epidermal  cells  are  affected.  The 
upper  and  older  cells  of  the  nucellus,  where  not  too  much  disin- 
tegrated, also  show  an  increase  in  size  over  those  of  younger 
nucelli,  which  accounts  in  part  for  the  elongation  of  this  organ 
as  it  keeps  pace  with  the  growth  of  the  sac. 

Methods. — After  trying  one  per  cent,  solution  of  osmic  acid, 
one-half  per  cent,  and  one  per  cent,  chromic  acid,  a  saturated 
solution  of  corrosive  sublimate  in  seventy  per  cent,  alcohol  and 
the  last  two  plus  a  small  addition  of  acetic  acid  to  prevent 
shrinkage,  the  corrosive  sublimate  with  acetic  acid  was  found 
to  give  best  results.  The  sections  thus  fixed  also  took  best  the 
stains  used. 

After  this  method  of  fixing,  the  tissues  were  washed  in  seventy 
per  cent,  alcohol  containing  iodine,  gradually  transferred  to  ab- 
solute alcohol,  imbedded  in  paraffine  through  xylol,  stained  and 
mounted  in  balsam  in  the  usual  way.  Alcohol  safranin  fol- 


Fink  :     CONTRIBUTION    TO    THE    LIFE-HISTORY    OF    RUMEX.     151 

lowed  by  methyl  blue  was  found  to  be  the  best  stain  for  stages 
before  the  establishment  of  the  macrospore,  and  safranin  or  Del- 
afield's  hasmatoxylin  gave  best  results  from  this  period  up  to  the 
fecundation  of  the  .egg,  after  which  the  hasmatoxylin  proved 
best. 

The  nucellus  of  Polygonwn  ereclum  is  transparent  enough  to 
show  the  nuclei  of  various  stages  of  development  of  the  sac,  ex- 
cept the  antipodals,  as  well  as  the  cytoplasm  and  vacuoles  with 
oil  immersion  lens,  without  sectioning  or  any  treatment  what- 
ever. The  results,  however,  were  not  reliable  enough  for  my 
purpose,  nor  are  they  certain  enough  for  use  in  instruction. 

The  figures  are  all  drawn  to  the  same  scale  by  using  one-inch 
Leitz  eye  piece,  one  and  one-half-inch  Leitz  objective  and 
camera  lucida. 


EXPLANATION  OF  PLATES. 

NOTE. — All  figures  are  of  Rumex  verticillatus  unless  otherwise 
indicated.  The  figures  of  the  first  two  plates  are  reduced  to  one-half 
the  original  size  of  drawings,  those  of  the  last  two  to  one-third. 

Plate  IX. 

1 .  Upper  portion  of  nucellus  showing  the  archesporium. 

2.  Later  stage  showing  the  primary  tapetum  cut  off  above  and  the 
mother  cell  of  the  embryo  sac  below. 

3.  The  tapetum  has  divided  into  two  tapetal  cells  and  the  mother 
cell  has  increased  considerably  in  size. 

4.  Stage  between  2  and  3  showing  the  tapetum  dividing. 

5.  Four  tapetal  cells  and  the  elongated  mother  cell. 

6.  The  mother  cell  dividing  and  one  tapetal  cell  above. 

7.  The  four  potential  macrospores  derived  from  the  mother  cell. 

8.  The  macrospore  and  a  highly  refractive  cytoplasmic  cap,  repre- 
senting an  almost  completely  absorbed  cell,  either  a  tapetal  cell  or  the 
upper  one  of  the  potential  macrospores. 

9.  The  nucleus  of  the  macrospore  dividing. 

10.  A  somewhat  older  stage  than  7  and  showing  the  upper  three 
potential  macrospores  partly  absorbed  by  the  lowest  one,  which  is  to 
become  the  macrospore. 

1 1 .  Embryo  sac  containing  two  nuclei  resulting  from  the  division  of 
the  nucleus  of  the  macrospore,  the  lower  one  being  somewhat  larger. 
The  highly  refractive  remains  of  four  nearly  absorbed  cells  of  the  nu- 
cellus are  also  shown. 


152  MINNESOTA    BOTANICAL    STUDIES. 

Plate  X. 

12.  Embryo  sac  of  Rumex  salicifolius  showing  the  two  nuclei  as 
above  and  the  lower  one  also  slightly  larger. 

13.  Embryo  sac  of  Rumex  salicifolius  showing  the  four  nuclei  de- 
rived from  division  of  two  corresponding  to  those  figured  in  n  and  12, 
and  also  showing  the  persistent  spindle  in  the  anterior  end  of  the  sac. 

14.  Embryo  sac  of  Rtimex  salicifolius  showing  the  corresponding 
four  nuclei  in  a  somewhat  older  sac. 

15.  Embryo  sac  of  Rzimex  salicifolius  showing  eight  nuclei  derived 
from  four  corresponding  to  those  figured  in   13   and  14.     The  polar 
nuclei  are  approaching  each  other.     The  three  anterior  nuclei  are  of 
about  equal  size  and  not  enclosed  in  walls.     Only  the  nucleoli  of  the 
antipodal s  could  be  seen. 

1 6.  Embryo  sac  of  Polygonum  erectum  showing  the  earlier  forma- 
tion of  cell  walls  about  the  anterior  nuclei,  one  of  which  was  lying  be- 
low the  other  two.     The  polar  nuclei  are  approaching  each  other,  and 
the  cellular  structure  in  the  antipodal  area  is  distinct,  one  cell  here  also 
lying  below  the  other  two. 

17.  Embryo  sac  of  Rumex  salicifolitts  showing  the  egg  yet  free, 
cell  walls  forming  about  the  synergidae,  the  polar  nuclei  fusing  and 
three  distinct  nucleated  antipodal  cells. 

18.  Embryo  sac  showing  the  sister  nuclei  which  form  the  synergidae 
yet  free  and  the  wall  forming  about  the  egg  after  the  definitive  nucleus 
is  formed. 

19.  Anterior  end  of  a  mature  embryo  sac  showing  the  synergidaa 
and  the  egg. 

20.  Embryo  sac  of  Rumex  salicifolius  showing  all  of  the  nuclei  of 
the  mature  sac,  except  the  antipodals  and  at  about  the   same  stage  as 
the  corresponding  nuclei  in  17. 

21.  Posterior  end  of  a  mature  embryo  sac  showing  three  antipodal 
cells. 

22.  Anterior  end  of  the  embryo  sac  of  Rumex  salicifolius  showing 
walls  forming  about  the  egg  after  the  definitive  nucleus  is  formed. 

23.  Mature  embryo  sac  showing  more  than  three  nuclei  in  the  anti- 
podal end. 

24.  Mature  embryo  sac  of  Rumex  salicifolius,  with  the  antipodal 
region  perhaps  not  distinctly  seen. 

Plate  XL 

25.  Anterior  end  of  embryo  sac  showing  one  synergid  degenerated, 
the   pollen  tube  entering  and  showing  two   nuclei  very  indistinctly. 
The  egg  and  definitive  nucleus  are  also  shown. 

26.  Anterior  end  of  embryo  sac  showing  a  pollen  tube  discharged 
and  containing  an  undischarged  sperm-nucleus.     The  two  synergidae 


VOL.    II. 


MINNESOTA    BOTANICAL    STUDIES.  PART    II. 


11 


PLATE    IX. 


VOL.    II. 


MINNESOTA    BOTANICAL    STUDIES.  PART    II. 


22 


17 


15 


!§ 


PLATE    X. 


HLLTCTlTr  PRTHTTWi;  •  J 


VOL.    II. 


MINNESOTA    BOTANICAL    STUDIES.  PART    II 


PLATE  XL 


VOL.    II. 


MINNESOTA    BOTANICAL    STUDIES. 


PLATE    X1T. 


Fink  :     CONTRIBUTION    TO    THE    LIFE-HISTORY    OF    RUMEX.      153 

have  degenerated,  the  fecundated  egg  is  extending  its  cell  wall  up- 
ward to  attach  it  to  the  anterior  end  of  the  sac  and  the  definitive  nu- 
cleus is  dividing. 

27.  Posterior  end  of  the  sac  whose  anterior  end  is  shown  in  26. 
The  antipodals  are  degenerating. 

28.  Embryo  sac  and    surrounding  tissue  of    the   nucleus  showing 
mutual  relations  of  the  two  structures. 

29.  Anterior  portion  of  embryo  sac  showing  three  of  a  probable 
four  endosperm  nuclei,  pollen  tube  ruptured  at  the  end  and  the  egg  di- 
viding to  form  the  two-celled  embryo. 

Plate  XII. 

30.  Anterior  end  of  an  embryo  sac  somewhat  younger  than  that  fig- 
ured in  29  and  showing  the  discharged  pollen  tube,  the  egg  apparently 
preparing  to  divide,  and  four  endosperm  nuclei  with  a  spindle  persist- 
ing between  each  pair. 

3 1 .  Anterior  end  of  embryo  sac  showing  a  five-celled  embryo  and 
three  endosperm  nuclei. 

32.  Young  embryo  in  anterior  end  of  embryo   sac  showing  four 
newly  formed  nuclei  and  two  persistent  spindles. 

33.  Embryo  sac  of  Rumex  salicifolius  showing  a  somewhat  more 
advanced  embryo,  a  larger  number  of  endosperm  nuclei  and  the  anti- 
podals still  persisting. 

34.  Anterior  end  of  embryo  sac  showing  five-celled  embryo,  one  of 
four  observed  dividing  endosperm  nuclei,  and  an  undischarged  pollen 
tube  which  entered  the  sac  after  the  egg  was  fecundated. 

35.  Anterior  end  of   embryo  sac  of  Rumex  salicifolius  showing 
three-celled  embryo,  whose  middle  cell  is  dividing,  and  the  persistent 
discharged  pollen  tube  in  the  micropyle  and  extending  into  the  sac. 


XL    OBSERVATIONS   ON  GIGARTINA. 


MARY  E.  OLSON. 


The  genus  Gigartina  is  of  wide  distribution,  especially  in 
temperate  latitudes.  It  is  found  both  in  the  Atlantic  and  Pa- 
cific oceans  as  far  north  as  the  coast  of  Greenland  and  as  far 
south  as  Cape  Horn. 

The  material  used  in  preparing  this  paper  was  collected 
in  Puget  Sound,  at  Channel  Rocks,  near  Seattle,  Washington, 
on  August  3,  1897.  It  does  not  correspond  exactly  with  any  of 
the  specific  descriptions  recorded.  Indeed  it  has  been  with 
some  reluctance  that  this  plant  has  been  included  in  Gigartina. 

The  material  used  for  study  in  the  preparation  of  this  paper 
had  been  preserved  in  75  per  cent,  alcohol,  with  the  exception 
of  some  dried  material  which  was  studied  in  determining  the 
color,  size,  shape  and  other  external  characters. 

In  the  dried  material  the  fronds  appear  thin  and  membrana- 
ceous  and  very  brittle.  The  alcoholic  material  is  leathery  in 
texture  and  quite  tough.  The  older  fronds  show  a  considerable 
increase  in  thickness  over  the  younger  ones,  and  are  strongly 
Calltblepharis-\\\iQ.  in  appearance. 

HABIT  AND  EXTERNAL  APPEARANCE. 

The  fronds  are  reddish  purple  in  color  and  occur  at  a  depth 
of  eight  fathoms.  Several  or  more  fronds  are  generally  found 
growing  crowded  together  from  united  holdfasts.  Such  a  group 
is  seen  in  Fig.  i,  PI.  13. 

The  general  outline  of  a  frond  whether  branched  or  un- 
branched  is  typically  cuneate.  Fig.  3,  PI.  13  shows  a  good 
specimen  of  the  unbranched  type.  When  branched  the  general 
wedge-shaped  outline  is  retained  by  a  more  or  less  regular, 
dichotomous  form  of  branching,  in  which  the  branches  spread 
and  remain  of  considerable  width.  (Fig.  i,  PL  13.) 


Olson  :     OBSERVATIONS    ON    GIGAKTINA.  155 

Holdfast. — The  holdfast  is  disc-like,  and  varies  in  size  accord- 
ing to  the  number  of  plants  springing  from  it.  In  an  isolated  in- 
dividual (Fig.  3,  PL  13)  it  is  seen  to  be  but  little  larger  than  the 
circumference  of  the  base  of  the  stipe.  On  the  under  side,  by 
which  it  is  attached  to  the  substratum,  it  is  seen  to  have  a  smooth 
surface,  and  two  areas  are  clearly  distinguishable  in  the  alco- 
holic material  used  in  this  study ;  a  central,  nearly  circular  al- 
most translucent  area,  and  an  outer  apparently  denser  portion 
(Fig.  3,  PI.  13).  When  viewed  from  the  upper  side  the  signi- 
ficance of  these  areas  is  understood.  If  a  large  holdfast  be  ex- 
amined, from  which  part  of  the  fronds  have  been  removed,  the 
upper  surface  will  be  seen  to  be  uneven  and  dotted  with  circu- 
lar pits  bordered  by  very  distinct  rims  (Fig.  6,  PI.  13).  On 
running  the  point  of  a  needle  through  one  of  the  translucent 
areas  of  the  under  side  it  is  found  to  correspond  with  the  pit  of 
the  upper  side.  A  still  more  interesting  demonstration  is  to  pull 
away  one  of  the  fronds  still  attached  to  the  holdfast.  If  this  be 
done  carefully  it  will  be  seen  that  a  characteristic  pit  remains  to 
mark  the  point  of  attachment  of  the  stipe.  An  old  holdfast  is 
found  to  be  covered  with  these  scars,  which  are  very  perfect 
markings  of  the  outline  of  the  stipe  at  the  point  of  attachment. 

Stipe. — The  stipe  is  a  well-marked  organ  whose  presence 
is  more  or  less  evident  in  all  the  plants,  especially  as  they  attain 
their  mature  size.  Some  of  the  members  of  the  group  in  Fig. 
i,  PL  13,  show  that  in  the  first  stages  the  width  of  the  frond 
varies  but  little  from  the  base  to  the  tip,  so  that  a  distinct  stipe 
is  scarcely  distinguishable.  Very  soon,  however,  the  upper  por- 
tion of  the  frond  begins  to  expand  and  a  typical  stipe  becomes 
evident.  Its  outline  just  at  the  point  of  attachment  to  the  hold- 
fast is  circular,  but  above  this  it  becomes  slightly  compressed  in 
one  diameter  so  that  its  cross  section  appears  oval  or  elliptical. 
The  transition  trom  stipe  to  lamina  is  so  gradual  that  no  distinct 
demarcation  can  be  detected  between  them.  The  shorter  diam- 
eter becomes  still  shorter,  and  the  longer  one  increases  to  the 
width  of  the  frond  till  all  appearance  of  the  stipe  has  van- 
ished and  even  the  greater  thickness  at  the  center  of  the  lamina 
merges  so  gradually  into  the  thinner  margins  that  it  cannot  be 
be  said  to  be  present  as  a  midrib. 

Lamina. — The  lamina  is  seen  to  attain  its  greatest  width  at 
some  little  distance  from  the  tip.  If  it  is  branched  the  division 
is  dichotomous,  though  the  lobes  are  often  unequal  in  size,  and 


156  MINNESOTA    BOTANICAL    STUDIES. 

occurs  near  the  tip  of  the  frond,  so  that  there  is  no  branching  of 
the  stipe.  The  margins  present  a  more  or  less  wavy,  undula- 
ting outline.  In  fertile  fronds  the  margins  all  along  the  upper 
part  of  the  frond  are  prolonged  into  little  leaf-like  outgrowths 
bearing  the  cystocarps  (Fig.  2,  PI.  13).  These  proliferations 
also  occur  on  the  surface  of  the  frond  and  sometimes  are  scarely 
more  than  the  stalk  of  the  single  cystocarps  they  bear.  In  many 
fertile  fronds  the  surface  is  almost  entirely  covered  with  these 
outgrowths.  As  a  rule  they  occur  much  more  densely  on  one 
side  than  the  other.  Frequently  they  are  found  as  simple  small 
leaflets  bearing  no  cystocarps.  Their  presence  gives  a  look  to 
the  lamina  quite  suggestive  of  fruiting  Calliblepharis. 

INTERNAL  ANATOMY. 

Holdfast. — A  section  of  the  holdfast  shows  a  very  distinctive 
structure  unlike  anything  seen  elsewhere  in  the  plant.  The  ap- 
pearance of  a  section  is  seen  in  Fig.  9,  PL  13,  showing  the 
outline  of  the  pit  or  scar  and  the  depression  from  which  the 
stipe  was  removed.  The  cellular  structure  varies  but  little  from 
the  general  type  except  in  the  transition  zone  from  the  holdfast 
to  the  stipe.  Along  the  upper  surface  the  cells  are  covered  by 
a  gelatinous  envelope  or  cuticle  of  considerable  thickness  often 
15  mic.  deep  (Fig.  10  a,  PL  13).  This,  of  course,  in  one  of  the 
scars  extends  only  to  the  border  of  the  pit  as  indicated  in  Fig.  9, 
PL  13.  Some  sections  also  show  a  similar  layer  on  the  lower  side 
at  joints  (Fig.  9  b,  PL  9),  while  the  remainder  of  the  lower 
surface  shows  a  rough  irregular  margin  of  cells  (Fig.  9  p,  PL 
13).  The  possible  explanation  suggests  itself  that  the  cuticle, 
when  occurring  on  the  lower  side,  appears  at  joints  which, 
through  some  unevenness  of  the  substratum,  are  not  closely  ap- 
pressed  to  it  and  hence  are  left  exposed  as  it  were. 

The  tissue  of  the  holdfast  is  characterized  by  cells  having  a 
quadrilateral  outline  as  seen  in  zone  "  b"  in  the  right-hand  part 
of  Fig.  10,  PL  13.  This  general  type  becomes  greatly  modi- 
fied in  various  regions.  In  the  transition  zone  "  c"  (Fig.  10, 
PL  13),  from  the  holdfast  to  the  stipe,  the  angular  outline  of  the 
cells  disappears  and  they  become  more  rounded  and  smaller  in 
size.  In  all  regions  cells  of  irregular  outline  are  frequently 
met  with  scattered  among  the  cells  of  typical  quadrilateral  out- 
line. 

In  general  the  cells  are  arranged  in  approximately  regular 


Olson  :     OBSERVATIONS    ON    GIGARTINA.  157 

rows  extending  vertically  through  the  holdfast.  This  is  more 
apparent  outside  the  region  of  the  pit  than  within.  In  passing 
from  the  holdfast  proper  to  the  stipe  the  character  of  the  cells  is 
seen  to  change,  the  outline  becomes  rounded,  the  cells  smaller 
and  the  arrangement  very  irregular  (Fig.  10  c,  PI.  13).  On 
the  upper  side  of  this  zone  the  cells  become  elongated  and  soon 
merge  into  the  structure  of  the  stipe  (Fig.  10  d,  PL  13).  At 
the  side,  in  the  region  where  the  surface  of  the  holdfast  passes 
to  the  surface  of  the  stipe,  an  interesting  curvature  of  the  rows  of 
cells  of  the  holdfast  is  noticed  (Fig.  10  e,  PI.  13).  The  upper 
cells  of  the  holdfast  gradually  become  smaller  and  merge  into 
the  outer  cells  of  the  stipe,  so  that  at  the  periphery  there  is  not 
so  marked  a  change  in  the  character  of  the  cells  as  in  the 
center.  The  central  elongated  cells  of  this  stipe  extend  farther 
into  the  holdfast  in  the  center  than  at  the  periphery  of  the  stipe 
region.  This  is  to  be  expected  from  the  outline  of  the  pits 
from  which  the  stipe  has  been  removed,  and  an  examination  of 
these  pits  under  high  power  reveals  the  fact  that  none  of  the 
elongated  stipe  cells  are  present  in  the  scar,  showing  that  the 
separation  zone  when  a  stipe  is  pulled  from  a  holdfast  is  at  the 
region  shown  in  Fig.  10,  PI.  13  at  c,  and  curves  upward  at  the 
sides,  thereby  forming  the  pit-like  scar. 

In  many  sections  of  stipe  and  holdfast  the  elongated  cells  are 
seen  to  extend  much  farther  down  than  in  the  section  of  Fig. 
10,  PI.  13,  so  that  only  one  or  two  layers  of  holdfast  cells  lie 
between  them  and  the  lower  surface. 

In  a  few  sections  an  interesting  development  of  outgrowths  on 
the  lower  surface  was  noticed.  These  occur  below  the  stipe 
region  and  show  the  same  cellular  structure  as  the  holdfast 
proper.  They  are  apparently  rhizoid-like  growths  (Fig.  10, 
PI.  13). 

The  sections  used  were  placed  in  an  alcoholic  solution  of 
fuchsin  for  a  few  moments,  then  washed  with  alcohol  and 
mounted  in  glycerine  jelly.  The  stain  failed  to  bring  out  any 
cell  contents  and  to  all  appearances  the  cells  are  empty. 

Stipe  and  Lamina. — The  structure  of  the  stipe  and  lamina  is 
very  similar,  the  chief  difference  being  that  in  the  former  the 
cells  are  of  somewhat  smaller  diameter. 

The  upright  portion  of  the  plant  may  be  divided,  anatomic- 
ally, into  two  fairly  distinct  portions:  the  pith,  consisting 
of  the  larger,  apparently  empty  cells  and  the  cortex,  of  smaller 


158  MINNESOTA    BOTANICAL    STUDIES. 

cells  containing  the  chromatophores.  Because  of  this  dif- 
ference in  the  cells  the  sections  for  the  structure  of  the  stipe 
and  lamina  show  the  outlines  of  the  cell  wall  in  the  pith,  but  in 
nearly  all  the  sections  examined  the  cell  walls  of  the  cortex 
could  not  be  distinguished,  and  it  is  the  outline  of  the  cell  con- 
tents that  is  represented  in  such  sections. 

The  inner  portion  or  pith  consists  of  elongated,  cylindrical 
cells,  united  into  loosely  interwoven  filaments,  extending  prin- 
cipally in  the  direction  of  the  long  axis  of  the  frond.  The 
union  between  cells  is  so  irregular  that  often  the  filamentous 
arrangement  is  scarcely  recognizable.  Surrounding  this  pith 
region,  which  is  of  compressed  cylindrical  contour  in  the  stipe, 
is  the  cortex,  which  consists  of  much  smaller  cells,  arranged  in 
radiating  rows,  more  or  less  regular,  perpendicular  to  the  sur 
face  of  the  frond. 

Pith. — The  transverse  sections  of  both  stipe  and  lamina  show 
the  cross-sectional  outline  of  the  pith  cells  to  be  more  or  less 
circular  with  considerable  space  between  the  cells  (Figs.  12  and 
13,  PL  13).  In  the  lamina  portions  of  a  filament  are  often 
found  running  through  the  section.  The  outlines  of  the  pith 
cells  in  both  lamina  and  stipe  show  an  area  of  larger  cells  just 
within  the  cortex,  passing  inward  to  a  central  portion  of  smaller 
diameter  (Fig.  n,  PL  13).  Within  the  pith  itself  there  is  con- 
siderable variation  in  the  size  of  the  cells,  showing  that  smaller 
filaments  anastomose  with  the  larger  ones. 

The  average  size  of  the  cells  is  from  100  to  170  mic.  long  by 
J7  to  33  mic.  wide.  The  most  interesting  feature  of  the  pith  is 
the  presence  of  protoplasmic  pits  connecting  the  cells.  These 
occur  not  only  in  the  end  walls,  but  also  in  the  lateral  walls,  as 
seen  in  Fig.  n,  PL  12.  By  these  connections,  as  well  as  by 
lateral  pressure  in  some  cases,  the  cylindrical  outline  of  the 
cells  becomes  variously  modified. 

Communication  of  adjacent  cells  of  the  pith  region  is  thus 
completely  established.  The  significance  of  this  will  be  more 
clearly  seen  as  the  physiological  importance  of  this  area  is  dis- 
cussed. These  pits  were  first  discovered  by  staining  with  haema- 
toxylin.  A  more  careful  trial  of  different  stains  showed  that 
these  pits  always  take  the  stain  more  deeply  than  any  other  part, 
either  cell  contents  or  cell  wall.  Both  methyl  violet  andfuchsin 
produced  good  results.  An  alcoholic  solution  of  the  stain  was 
used. 


Olson  :    OBSERVATIONS    ON    GIGARTINA.  159 

A  study  of  these  pits  showed  them  to  be  of  the  form  of  small 
plates  or  rings,  apparently  one  in  each  of  the  two  adjacent  cells 
(Figs,  ii  and  13,  PI.  9).  When  seen  edgewise  they  appear  as 
two  small  plates  narrower  than  the  width  of  the  cell  wall,  so  that 
the  inner  line  of  the  cell  wall  appears  to  curve  out  to  meet  them. 
Often  the  entire  outline  of  one  ring  may  be  seen  .and  only  part 
of  the  other,  which  apparently  lies  beneath  it.  Again  "the  sec- 
tion will  lie  so  as  to  show  both  rings. 

Some  sections,  especially  with  the  methyl  violet,  showed  a 
faint  outline  of  cell  contents  just  within  the  wall  and  in  all  cases 
extending  close  up  to  the  rings  or  pits.  Careful  observations  of 
this  sort  led  to  the  opinion  that  these  connecting  pits  uiust  be  of 
the  nature  of  the  protoplasmic  cell  contents  rather  than  the  cell 
wall.  Schmitz  confirms  this  opinion.  Unfortunately  the  writer 
did  not  have  access  to  Schmitz's  original  article,  but  in  George 
Murray's  Introduction  to  the  study  of  Seaweeds,  1895,  in  the 
chapter  on  Rhodophycese  (which  he  states  is  based  upon 
Schmitz's  papers)  the  following  is  found  (p.  201):  "The 
plates  stand  in  direct  connection  with  the  protoplasm  lining  the 
cell  wall  and  are,  in  fact,  so  coherent  with  it  that  they  may  be 
regarded  as  transformed  or  rather  differentiated  protoplasm  lo- 
cally covering  the  pit.  However  it  is  probable  that  a  thin  layer 
of  protoplasm  covers  them  in  turn." 

All  observations  have  gone  to  show  that  there  is  an  intimate 
protoplasmic  connection  between  the  contents  of  neighboring 
cells.  Zimmermann's  Botanical  Microtechnique  was  consulted 
as  to  re-agents  for  testing  these  rings. 

The  use  of  sulphuric  acid  and  a  mixture  of  iodine  and  potas- 
sium iodide  is  recommended  for  cellulose  walls  giving  a  blue 
color.  This  was  used,  but  neither  the  cell  walls  nor  the  pits 
showed  any  trace  of  blue  staining.  Cuprammonia  was  also 
tried,  but  with  no  success.  Some  interesting  results  were  ob- 
tained, however,  with  the  use  of  sulphuric  acid.  The  sections 
were  first  stained  and  then  treated  with  the  acid.  Although  the 
acid  at  once  destroyed  the  original  color  it  was  found  better  re- 
sults were  obtained  than  without  first  staining.  A  trial  was  then 
made  as  to  the  strength  of  acid  which  would  give  most  satisfac- 
tory results,  and  it  was  found  that  treating  sections  prepared 
as  before  described,  with  a  50  to  60  per  cent,  solution  of  the  acid 
produces  at  first  no  apparent  effect  beyond  a  slight  swelling  of 
the  cell  wall.  The  sections  were  left  mounted  in  the  acid,  and 


160  MINNESOTA    BOTANICAL    STUDIES. 

after  twenty-four  hours  re-examined.  It  was  then  found  that 
the  cell  walls  were  all  dissolved  and  only  the  rings  remained 
(Fig.  17,  PL  14).  This  leads  to  the  conclusion  that  the  rings 
are  not  of  the  same  composition  as  the  cell  wall. 

The  pith  of  the  cystocarpic  proliferations  of  the  frond  shows 
a  marked  difference  from  that  of  the  vegetative  portion  (Fig. 
18,  PL  14).  The  cells  have  become  very  irregular  in  outline 
and  are  so  anastomosed  and  interwoven  as  to  form  a  network 
which  becomes  more  and  more  dense  in  passing  from  the  stalk 
to  the  pericarp  proper.  Here,  as  well  as  in  the  cortex,  the  cell 
contents  have  a  dense  granular  appearance,  and  the  cell  walls 
appear  only  very  faintly,  if  at  all.  In  most  sections  stained  as 
in  the  vegetative  part  they  cannot  be  distinguished.  This  is  true 
also  of  the  protoplasmic  pits,  though  it  is  evident  there  is  close 
communication  throughout.  From  one  or  two  unusually  clear 
sections  it  was  ascertained,  however,  that  the  rings  are  present, 
but  are  very  small.  The  cells  in  this  region  measure  from  25 
to  37  mic.  long  by  2.5  to  7  mic.  wide. 

Cortex. — The  transition  from  the  pith  to  the  cortex  is  some- 
what abrupt.  In  longitudinal  sections  the  pith  cells  are  seen  to 
decrease  in  length  until  in  the  four  or  five  outermost  rows  the 
outline  of  the  cells  is  spherical  or  slightly  oblong.  In  the  transi- 
tion zone;  or  the  inner  part  of  the  cortex,  they  measure  from 
5  to  12  mic.  along  either  diameter. 

In  the  outermost  layers  of  smallest  cells,  measuring  from  2.5 
to  5  mic.  in  diameter,  the  cell  contents  are  very  dense  and  the 
cells  are  apparently  imbedded  in  a  gelatinous  matrix  from  which 
it  is  impossible  to  distinguish  their  walls.  In  one  section,  how- 
ever, the  writer  was  able  to  make  out  faint  outlines  of  the  walls, 
but  it  is  difficult  to  represent  them  and  maintain  the  proportional 
thickness  of  the  wall  (Fig.  n,  PL  13). 

The  cortex  cells  are  seen  to  lie  in  communication  also,  but 
only  along  the  radial  lines  of  the  thallus.  The  cells  are  so 
small  no  rings  can  be  distinguished,  but  protoplasmic  threads 
are  seen  running  from  cell  to  cell  (Fig.  13,  PL  13).  There  are 
no  lateral  protoplasmic  connections  between  cells.  A  surface 
view  of  the  thallus  shows  a  somewhat  regular  arrangement  of 
the  end  cells  of  these  radial  rows.  They  appear  as  a  rule  in 
groups  of  two  or  occasionally  three,  surrounded  by  the  gelatin- 
ous matrix  (Fig.  14,  PL  14).  If  the  sections  be  placed  in  water 
this  swells  rapidly,  as  do  also  the  cell  walls.  The  walls  often 


Olson:     OBSERVATIONS    ON    GIGARTINA.  161 

increase  to  three  or  four  times  their  width,  as  seen  in  alcoholic 
material,  and  a  stratification  of  the  walls  becomes  evident  (Fig. 
16,  PI.  14). 

REPRODUCTIVE  ORGANS. 

Cystocarps. — The  material  studied  was  too  far  advanced  to 
show  antheridia  or  the  development  of  the  cystocarp. 

The  cystocarps  are  found  scattered  very  abundantly  along  the 
margin  and  over  the  surfaces  of  the  fertile  fronds.  As  a  rule 
they  are  much  more  abundant  upon  one  surface  than  the  other. 

They  are  borne  in  leaf-like  proliferations  of  the  frond  and 
are  usually  more  or  less  distinctly  stalked,  though  often  they  are 
nearly  sessile.  Fig.  4  shows  one  of  these  leaf-like  outgrowths 
bearing  no  less  than  nine  cystocarps.  Generally  the  number  is 
smaller,  from  two  to  four  (Fig.  5,  PI.  13).  The  larger  leaflets, 
with  a  larger  number  of  cystocarps,  are  usually  found  along  the 
margin. 

In  form  the  cystocarps  are  subglobose  with  a  marked  indenta- 
tion at  the  apex  which  seems  to  indicate  a  distinct  carpostome, 
but  in  the  large  number  of  sections  observed  no  opening  could 
be  detected.  The  nearest  approach  to  it  was  seen  in  the  section 
represented  in  Fig.  19,  PI.  10,  but  even  here  there  is  no  sign  of 
a  true  pore  or  even  of  a  rupture,  so  that  evidently  the  cystocarp 
is  closed.  From  the  uniform  closure  of  the  cystocarp  it  is  diffi- 
cult to  include  the  plant  in  question  with  Gigartina.  The  struc- 
ture of  the  cystocarp  regions  of  the  thallus  shows  the  same  two 
general  areas  described  for  the  stipe  and  lamina  with  the  modi- 
fication of  the  inner  region  or  pith  described  above.  The  spores 
are  developed  within  the  central  region,  the  cortex  and  outer  part 
of  the  pith  forming  a  true  pericarp.  Except  at  the  apex  both 
areas  surround  the  central  mass  of  spores.  Here  it  is  covered 
only  by  the  cortex. 

A  peculiar  structure  observed  was  one  in  which  the  surface  of 
the  thallus  was  only  slightly  raised  to  indicate  its  location,  and 
numerous  long  filaments  were  seen  with  their  tips  protruding 
slightly  from  the  surface  (Fig.  18,  PI.  14).  The  section  was 
stained  with  fuchsin  and  the  clear  filaments  were  sharply  dis- 
tinguishable from  the  other  cells  with  their  granular  contents. 
They  measured  150  mic.  in  length. 

There  was  some  little  doubt  as  to  what  should  be  the  interpreta- 
tion of  the  section  represented  in  Fig.  18,  PI.  14.  The  pith  cells 


162  MINNESOTA    BOTANICAL    STUDIES. 

have  the  appearance  characteristic  of  the  cystocarp  region.  A 
comparison  with  figures  in  which  trichogynes  are  represented 
shows  but  slight  similarity  in  appearance.  The  pericarp  consists 
of  the  two  layers  found  in  the  lamina  and  stipe.  The  outer  small 
cells  containing  chromatophores  pass  somewhat  abruptly  to  the 
cells  of  the  interior  (Fig.  21,  PL  14),  which  are  elongated  and 
connected  to  form  a  more  or  less  dense  net-work.  In  most  cases 
the  transverse  connecting  cells  are  more  numerous  than  in  Fig. 
21,  so  that  the  pericarp  presents  the  reticulated  appearance  of 
the  tissue  in  Fig.  19.  Very  frequently,  however,  the  cells  show 
much  lateral  crowding  in  the  pericarp. 

The  cystocarp  is  compound  and  the  spores  are  aggregated 
into  distinct  groups  (Fig.  19,  PI.  14).  This  is  clearly  seen  in 
all  but  the  oldest  cystocarps  and  even  here  a  carefully  cut  sec- 
tion shows  it.  These  groups  are  separated  from  each  other  by 
large,  empty  cells,  with  smaller  cells  of  the  same  character  ex- 
tending between  them  (Fig.  20,  PI.  14).  This  is  brought  out 
very  clearly  by  staining  the  section  with  iodine  and  then  wash- 
ing in  water. 

The  carpospores  are  more  or  less  oval  in  shape,  often  some- 
what angular.  They  measure  from  12  to  15  mic.  along  one 
diameter  by  10  to  12  mic.  along  the  other. 

The  normal  cystocarps  measure  from  i  to  2  mm.  in  diameter, 
but  it  was  noticed  that  frequently  some  were  met  with  from  two 
to  three  times  as  large  as  the  ordinary  ones,  measuring  2.5  to 
3.5  mm.  On  examination  it  was  discovered  that  the  fronds  of 
another  small  alga  were  always  found  upon  these  large  cysto- 
carps. Several  specimens  were  studied  and  it  was  found  that 
there  was  evidently  more  than  one  species  infecting  them.  The 
largest  one  discovered  is  represented  in  Fig.  7  a,  PI.  13.  It  ap- 
pears to  emerge  directly  from  the  apical  depression  of  the  cys- 
tocarp. A  longitudinal  section  through  the  cystocarp  shows 
the  parasite  or  epiphyte  to  consist  of  an  axial  cylinder  of  large 
cells  with  protoplasmic  connections  between  adjacent  walls. 
Most  externally  is  a  region  of  quadrilateral  cells  larger  than  the 
corresponding  cells  of  the  host  arranged  quite  irregularly,  and  be- 
tween this  and  the  central  cylinder  a  region  of  long  filamentous 
cells,  and  it  is  these  which  are  seen  to  penetrate  the  pith  of  the 
pericarp  (Fig.  22,  PI.  14).  They  can  be  traced  to  the  central 
spore-bearing  area,  where  they  apparently  curve  outward  and 
follow  along  the  side  a  short  distance.  They  are  distinguish- 


Olson  :     OBSERVATIONS    ON    GIGARTINA.  163 

able  from  the  granular  cells  of  the  pericarp  by  their  clear  ap- 
pearance. 

In  most  cases  the  infecting  plant  was  found  to  be  a  smaller 
one,  represented  in  b  Fig.  7,  PL  13.  It  consists  of  simple  or 
branched  filaments  of  oblong  cells,  but  its  entrance  into  the  tis- 
sues of  the  pericarp  could  not  be  detected.  Its  presence  is  evi- 
dently the  cause  of  the  enlargement  of  the  cystocarp,  however. 
Except  in  cases  where  the  parasite  can  be  seen  within  the  tis- 
sues of  the  cystocarp  no  difference  except  size  can  be  observed 
between  the  infested  organs  and  the  normal  ones. 

Nemathecia. — These  were  found  upon  only  one  frond  in  the 
alcoholic  material  at  command,  but  in  this  they  were  abundantly 
distributed  on  both  sides  of  the  frond.  They  appear  in  surface 
view  as  wart-like  projections  which  can  be  distinguished  with 
the  naked  eye  by  their  slightly  lighter  color  (Fig.  8,  PI.  13). 
(It  must  be  remembered  that  this  description  applies  to  alcoholic 
material.) 

A  section  of  the  frond  shows  many  interesting  features.  The 
filaments  of  the  internal  pith  area  are  even  more  loosely  anas- 
tomosed than  in  the  vegetative  part  of  the  frond  and  show  large 
intercellular  spaces.  In  this  central  area  large,  dark  bodies  the 
size  of  spores  were  discovered  scattered  very  abundantly  among 
the  filaments  (Fig.  24  e,  PI.  14).  These  could  easily  be  seen 
from  the  surface,  showing  through  the  external  area  of  cortex 
cells,  and  even  appear  to  the  naked  eye  as  tiny  black  dots. 
Upon  examination  they  were  found  to  be  of  a  dark  green  color 
and  apparently  unicellular.  They  are  evidently  internal  para- 
sites, but  no  connection  between  them  and  the  nemathecia  could 
be  discovered,  though  it  was  earnestly  sought,  inasmuch  as 
Schmitz,  in  his  article  "Die  Gattung  Actinococcus  Kutz,"* 
ascribes  the  nemathecia  of  Phyllo-phora  brodi&i  and  P.  inter- 
rupta  to  the  parasite  Actinococcus. 

Outside  the  central  filaments  is  an  area  of  approximately 
spherical  cells,  which  decrease  gradually  in  size  toward  the  ex- 
terior. They  exhibit  a  characteristic  arrangement,  i.  e.,  from  a 
single  basal  cell  two  and  sometimes  three  rows  diverge  toward 
the  surface.  The  outer  layer  is  covered  by  a  thick  gelatinous 
cuticle  (Fig.  24,  PI.  14).  The  tetraspores  are  evidently  pro- 
duced from  the  cells  just  outside  the  central  filaments  and  are 
formed  in  irregular  masses  just  below  the  surface. 

*  Schmitz.     Flora.  77  :  367.     1893. 


164  MINNESOTA    BOTANICAL    STUDIES. 

In  many  places  where  there  is  almost  no  elevation  of  the  sur- 
face a  small  group  of  tetraspores  is  distinctly  seen  in  section. 
Staining  with  iodine  brings  out  the  distinction  between  tetras- 
pores and  the  surrounding  cells  very  clearly. 

A  peculiar  feature  of  the  nemathecia  is  the  pore-like  break  in 
the  cuticle  just  above  the  group  of  spores.  This  was  seen  in 
even  the  smallest  nemathecia,  and  in  the  larger  ones  several 
were  often  present.  A  full-grown  nemathecium  rises  from  25-37 
mic.  above  the  level  of  the  thallus,  and  an  irregularity  in  the  ar- 
rangement of  the  cortex  cells  is  noticeable  at  the  apex,  suggest- 
ing a  rupture  as  a  result  of  the  crowding  upward  of  the  spores. 
The  spores  measure  12-20  mic.  by  10-15  mic. 

Wille*  makes  a  physiological  distinction  between  the  pith  and 
cortex  regions,  considering  the  former  as  conducting  tissue  and 
the  latter  as  assimilative.  Some  interesting  results  were  brought 
out  by  iodine  staining  in  connection  with  this  view. 

Sections  of  the  holdfast,  stipe,  lamina,  cystocarp  region  and 
nemathecia  were  placed  in  an  iodine  solution  for  half  an  hour, 
then  washed  with  water,  with  the  following  results  :  The  hold- 
fast simply  shows  a  general  yellowish  staining  of  the  cell  walls, 
showing  there  is  no  starch  present  in  that  region,  as  would  be 
expected  from  its  evident  mechanical  function.  The  stipe  like- 
wise showed  a  slight  yellowish  staining  of  the  walls,  but  no  cell 
contents  in  accordance  with  its  function  as  a  supporting  and 
conducting  area. 

In  the  lamina  the  sections  used  were  longitudinal  ones.  The 
central  elongated  cells  remained  colorless ;  in  the  inner  cortex 
cells  marking  the  transition  in  shape  from  the  central  to  the 
small  peripheral  cells  the  contents  stained  a  deep  purple,  in- 
dicating the  presence  of  starch.  The  four  or  five  outer  rows 
showed  the  cell  contents  stained  yellowish  brown.  These  are 
the  cells  containing  chromatophores.  These  results  suggest  a 
confirmation  of  Miiller's  interpretation  of  the  physiological  sig- 
nificance of  this  area,  inasmuch  as  they  are  evidently  concerned 
with  the  food  supply. 

In  the  cystocarp  region  the  outer  layers  of  cells  containing 
rhodoplasts  stain  yellowish  brown  as  in  the  lamina ;  the  rest  of 
the  pericarp  and  the  spores  stain  a  deep  violet,  but  the  thin- 
walled  cells  separating  the  groups  of  spores  stain  yellow.  This 
brings  out  the  structure  of  the  central  spore  region  better  than 
any  of  the  other  stains  used. 

*  Wille,  N.   Nova  Acta  Acad.  Leop. -Carol.  Nat.  Cur.  52  :  49-100.  pi.  3-8.    1897. 


Olson  :     OBSERVATIONS    ON    GIGART1NA.  165 

In  the  nemathecia  the  tetraspores  stain  a  deep  violet,  as  also 
do  the  contents  of  the  central  filaments  just  below  the  spores. 
The  rest  of  the  central  filaments  remain  unaffected  and  the  outer 
cells  stain  yellowish  brown. 

SPECIFIC  DESCRIPTION. 

Gigartina  sp.  und. — Fronds  purplish-red,  distinctly  caules- 
cent, several  often  springing  from  the  same  disc-like  holdfast ; 
18-28  cm.  long  by  7-10  cm.  wide.  Stipe  somewhat  compressed, 
3.5  to  5  mm.  wide  by  2-3  mm.  thick,  gradually  widening  into 
the  typically  cuneate  lamina.  Young  fronds  often  entire,  older 
ones  sparingly  branched,  branches  expanded,  never  linear  or 
lanceolate.  Cystocarps  compound,  closed,  more  or  less  stalked, 
several  generally  occurring  crowded  together  on  the  same  pro- 
liferation ;  enclosed  within  a  pericarp;  1-2.5  rnm.  in  diameter; 
carpospores  numerous,  crowded  together  in  more  or  less  definite 
groups,  oval,  12-15  mic.  long  by  10-12  mic.  wide.  Tetraspores 
produced  in  nemathecia,  on  both  sides  of  the  frond ;  nemathecia 
wart-like,  rising  25-37  mic.  above  the  level  of  the  thallus.  Tet- 
raspores oval,  more  or  less  angular,  12-20  mic.  long  by  5~10 
mic.  wide. 

METHODS. 

Part  of  the  sections  used  were  cut  with  the  freezing  microtome, 
the  rest  by  hand.  The  material  used  had  been  preserved  in  75 
per  cent,  alcohol.  The  effect  of  imbedding  in  gelatine  pre- 
pared according  to  Osterhout's  directions  was  tested.  The  por- 
tions to  be  sectioned  were  cut  in*  pieces  .5  cm.  long  and  about 
the  same  width  and  placed  in  the  gelatine.  This  was  left  for 
twenty-four  hours  to  allow  the  gelatine  to  penetrate  the  tissues, 
then  removed  and  placed  in  a  gum  arabic  solution  on  the  freez- 
ing chamber.  After  several  trials  it  was  found  that  sections  in- 
troduced directly  into  the  gum  arabic  without  embedding  were 
as  satisfactory  as  by  the  longer  process. 

Staining. — At  first  water  solutions  of  the  stains  were  used, 
but  it  was  found  that  this  caused  the  tissues  to  swell  to  such  an 
extent  that  cells  often  presented  a  very  unnatural  appearance. 
For  example,  a  cross  section  of  the  stipe  was  obtained  in  which 
the  cell  lumen  appeared  irregulary  star-shaped  or  was  nearly 
obliterated  (Fig.  16,  PI.  14).  This  was  then  abandoned  and 

*Osterhout.     Bot.  Gaz.  21:   195-201.    1896. 


166  MINNESOTA    BOTANICAL    STUDIES. 

alcoholic  solutions  were  used.  Both  staining  the  sections  and 
the  material  in  toto  were  tried.  In  the  latter  method,  if  the  ma- 
terial was  to  be  imbedded,  it  was  first  stained.  If  the  stain  is 
sufficiently  diluted,  and  the  sections  are  allowed  to  stand  in  it 
from  ten  minutes  to  half  an  hour,  the  results  are  quite  as  satis- 
factory as  staining  in  toto,  for  in  the  latter  method  often  the  stain 
fails  to  penetrate  the  material  completely. 

The  stains  employed  were  hasmatoxylin,  anilin  blue,  methyl 
blue,  carmine,  methyl  violet,  fuchsin  and  safranin.  Hasma- 
toxylin  stains  both  cell  wall  and  contents,  but  not  clearly 
enough ;  it  was  found  very  unsatisfactory.  Anilin  blue  and 
carmine  hardly  affected  the  tissues.  Methyl  blue  and  safranin 
proved  good  for  the  gelatinous  sheath.  Methyl  violet  and 
fuchsin  gave  the  most  satisfactory  results.  They  stain  both 
the  cell  walls  and  protoplasmic  contents,  but  the  latter  more 
deeply.  These  two  stains,  and  especially  fuchsin,  were  used 
for  all  the  work. 

The  sections  were  at  first  taken  from  the  knife  and  placed  in 
glycerine,  transferring  gradually  from  20  per  cent,  to  absolute, 
but  the  glycerine  was  found  to  swell  the  cell  walls  to  a  consider- 
able extent  and  was  abandoned. 

The  most  satisfactory  method  and  the  one  employed  in  all  the 
latter  part  of  the  work  was  as  follows  :  The  alcoholic  material 
was  placed  directly  in  the  gum  arabic  solution  on  the  freezing 
chamber,  transferred  from  the  knife  to  the  alcoholic  solution  of 
the  stain  and  mounted  in  it.  When  the  staining  was  completed, 
usually  after  a  few  moments,  the  sections  were  washed  in  alco- 
hol. This  was  then  evaporated  and,  without  allowing  the  sec- 
tions to  become  dry,  glycerine  jelly  was  added,  making  a  per- 
manent mount. 

BIBLIOGRAPHY. 

Jonsson,  B.  Beitrage  zur  Kenntniss  des  Dickenzuwachses  der 
Rhodophyceen.  (Bot.  Zeit.  50:  181.  1892.)  (Bot.  Jahresb.  19: 
120— 121.  1894.)  Lunds  Univers.  Arskr.  27:  41.  2  pi.  1890—91. 

Wille,  N.  Beitrag  zur  Entwicklungsgeschichte  der  physiologischen 
Gewebesysteme  bei  einigen  Florideen.  Nova  Acta  Acad.  Leop.- 
Carol.  52:  49-100.  pi.  3-8.  1887.  (Bot.  Jahresb.  16:  754. 
1891.)  " 

Osterhout,  W.  J.  V.  A  Simple  Freezing  Device.  Bot.  Gaz.  21  : 
195-201.  1896. 

Schmitz,  F.  Die  Gattung  Actinococcus  Kiitz.  Flora.  77  :  367- 
418.  pi.  7.  1893. 


Olson:     OBSERVATIONS    ON    G1GARTINA.  167 

Moore,  S.  Le  M.  Studies  in  Vegetable  Biology.  —  I.  Observations 
on  the  Continuity  of  Protoplasm.  Journ.  Linn.  Soc.  21  :  595-621.  pt. 
19-21.  1886. 

EXPLANATION  OF  FIGURES  IN  PLATES  XIII.  AND  XIV. 

All  drawings  were  made  from  material  preserved  in  alcohol. 

Fig.  i.  Group  of  fronds  springing  from  a  common  holdfast.  One- 
half  natural  size. 

Fig.  2.  Upper  portion  of  fertile  frond  showing  marginal  and  sur- 
face proliferations  with  cystocarps  produced  upon  them.  One-half 
natural  size. 

Fig.  3.  Young  frond  showing  typical  shape  and  single    holdfast. 

One-half  natural  size. 

Fig.  4.   Small  leaflet  with  cystocarps.     X4-5. 

Fig.  5.  Four  cystocarps  on  a  small  stalk-like  proliferation.     X4-5- 

Fig.  6.   Surface  view  of  holdfast  showing  scars.     X4-5- 

Fig.  7.  Cystocarps  infected  by  a  parasite.     x5. 

Fig.  8.   Surface  view  of  nemathecia.     x5- 

Fig.  9.  Section  through  a  scar  of  the  holdfast  :  #,  layer  of  cuticle 
on  upper  surface  ;  b,  region  of  the  lower  surface  covered  with  cuticle  ; 
c,  rough  broken  lower  surface,  x  5. 

Fig.  10.  Section  through  the  holdfast  with  rhizoid-like  outgrowth 
from  the  lower  surface.  A  portion  of  the  tissue  is  omitted  in  the  cen- 
ter, as  it  issimply  a  continuation  of  that  represented  on  either  side. 
Toward  the  left  the  drawing  stops  in  the  stipe  region.  On  the  right 
the  transition  from  the  stipe  to  the  holdfast  is  shown  :  a,  cuticle  of 
upper  surface;  6,  region  showing  cellular  structure  characteristic  of 
the  holdfast;  c,  cellular  structure  marking  transition  from  tissue  of 
holdfast  to  that  of  stipe  ;  d,  beginning  of  tissue  of  the  stipe  ;  e,  curva- 
ture of  cells  at  juncture  of  stipe  and  holdfast,  x  300. 

Fig.  1  1  .  Longitudinal  section  of  the  lamina  :  #,  cortex  ;  3,  outer 
region  of  pith  showing  lai-ger  pith  cells;  c,  inner  region  of  pith  with 
smaller  cells,  x  300. 

Fig.  12.  Cross  section  of  stipe:  «,  cortex;  £,  larger  celled  pith; 
c,  smaller  celled  pith,  x  300. 

Fig.  13.  Cross  section  of  the  lamina  showing  many  pith  cells  con- 
nected by  pits  and  cortex  cells  connected  by  protoplasmic  threads. 


Fig.  14.   Surface  view  of  thallus.     x  300. 

Fig.  15.  Outline  of  cross  section  of  stipe  showing  cortex  and  pith 
areas,  x  4.5. 

Fig.  1  6.  Pith  cells  of  lamina  stained  with  methyl  violet  and 
mounted  in  glycerine.  Cell  wall  is  much  swollen  and  one  shows 
stratification,  x  300. 


168  MINNESOTA    BOTANICAL    STUDIES. 

Fig.  17.  Effect  of  sulphuric  acid  on  cell  wall  and  pits;  a,  pith 
cells  treated  for  ten  to  fifteen  minutes  in  50  per  cent,  solution ;  b, 
pits  as  seen  twenty-four  hours  after  treating  the  section  with  the  acid. 
The  walls  have  been  entirely  dissolved,  x  300. 

Fig.  1 8.  Cluster  of  filaments,     x  300. 

Fig.  19.   Longitudinal  section  through  a  cystocarp.      x  85. 

Fig.  20.  Group  of  spores  from  cystocarp  with  thin-walled  cells 
separating  them,  x  300. 

Fig.  21 .  Longitudinal  section  through  the  pericarp  :  #,  cortex;  b, 
pith  ;  <:,  spores,  x  300. 

Fig.  22.  Longitudinal  section  through  an  infested  cystocarp.  The 
upper  left-hand  portion  sho\vs  the  manner  in  which  the  parasite  pene- 
trates the  host,  x  56. 

Fig.  23.  Portion  of  the  same  region  enlarged  showing  to  the  left 
the  tissues  of  the  host  pericarp  and  to  the  right  the  long  filamentous 
cells  of  the  parasite,  x  300. 

Fig.  24.  Section  through  frond  producing  nemathecia.  On  the 
upper  surface  is  a  mature  nemathecium,  on  the  lower  surface  two 
two  younger  ones  :  <z,  gelatinous  cuticle  ;  b.  pore-like  break  in  cuticle  ; 
c,  cortex  cells;  d^  cells  from  which  tetraspores  are  produced;  e,  para- 
site ;  f,  pith  filaments,  x  300. 


/OL.    II. 


MINNESOTA 


PL 


CAL    STUDIES. 


PART 


VOL.   II. 


MINNESOTA 


18 


14 


15 


16 


19 


;AL  STUDIES. 


PART 


21 


22 


24 


XII.  SEED    DISSEMINATION    AND     DISTRIBUTION 
OF  RAZOUMOFSKYA  ROBUSTA  (Engelm.)  Kuntze.* 


D.  T.   MACDOUGAL. 


The  branches  of  the  bull  pine  (Pt'nus  ponderosa  scopulorum), 
of  the  southwestern  United  States,  offer  suitable  conditions  of 
nourishment  for  the  growth  of  Razoumofskya  robusta,  a  parasite 
belonging  to  the  Loranthaceas.  Some  of  the  members  of  this 
family,  such  as  the  mistletoe  (Phoradendron  flavescens},  which 
live  on  deciduous  trees  in  temperate  latitudes,  are  furnished  with 
a  fair  amount  of  chlorophyll.  These  forms  are  able  to  carry  on 
more  or  less  food-formation  during  the  warmer  portions  of  the 
season  in  which  the  deciduous  hosts  lack  leaves.  Razoumofskya, 
however,  fastens  on  an  evergreen  conifer,  and  hence  has  no  such 
need  or  use  for  chlorophyll.  It  is,  therefore,  furnished  with  this 
substance  in  minute  quantity  only,  and  its  leaves  are  reduced  to 
mere  bracts.  It  is  dioecious,  and  the  aerial  shoots  of  both  kinds 
may  appear  in  close  contiguity  on  the  same  branch  of  the  host 
or  be  separated  some  distance.  The  shoots  start  up  from  the 
submerged  rhizomes  in  the  latter  part  of  April  or  early  in  May, 
the  flowers  maturing  in  June  and  the  seeds  in  August.  After 
the  dispersal  of  the  seeds  the  aerial  portion  of  the  plant  dies  away, 
leaving  only  the  haustorial  rhizomes  buried  in  the  tissues  of  the 
host  plant.  With  the  opening  of  the  next  season  shoots  are 
produced  as  before. 

The  submerged  portion  of  the  parasite  penetrates  the  branches 
of  the  host  long  distances  longitudinally,  and  where  aerial  shoots 
are  given  off  the  tissues  of  the  host  show  abnormal  structures, 
the  branches  undergoing  enlargement,  while  the  development 
of  the  nearest  buds  is  variously  checked  and  altered.  The  dis- 
tortion is  magnified  with  age,  and  old  trees  exhibit  the  most 
grotesque  malformations.  The  writer  has  seen  trees  a  meter  in 
height  infected,  and  the  size  of  the  older  branches  bearing  the 

*An  abstract  of  this  paper  was  read  before  the  Botanical  Club  of  the  A.A.A.S. 
at  Boston,  August  25,  1898. 


170  MINNESOTA    BOTANICAL    STUDIES. 

parasite  is  such  as  to  justify  the  statement  that  they  may  live  ten 
to  twenty  years  after  the  parasite  has  fastened  upon  them.  A 
common  type  of  structure  resulting  from  the  attachment  of  the 
parasite  to  the  pine  consists  of  an  old  branch  bent  downward, 
from  the  infected  tip  of  which  numbers  of  smaller  branches  stand 
erect,  forming  a  coarse  "  witch's  broom"  (See  Plate  XVI.). 

Perhaps  the  most  interesting  facts  in  connnection  with  the 
history  of  Razoumofskya,  are  tho.se  which  concern  the  distribu- 
tion of  the  seeds.  The  single-seeded  berries  are  borne  on  short 
stalks  curved  semi-circularly,  from  which  they  are  easily  de- 
tached when  ripe.  The  berry  is  joined  to  the  stalk  by  a  scission 
layer,  which  is  ruptured  by  the  slightest  touch  or  may  be  burst 
away  by  the  action  of  forces  set  up  in  the  berry,  which  also 
expel  the  seed.  The  shooting  of  the  seeds  from  the  berry  has 
been  known  for  many  years,  and  a  note  of  the  fact  has  found 
place  in  American  text-books  of  systematic  botany,  but  it  has 
failed  of  wider  recognition.  Engler  and  Prantl  remark  con- 
cerning the  seeds  of  the  Loranthaceae :  "  The  stickiness  enables 
some  seeds  falling  from  branch  to  branch  to  become  attached  ; 
on  the  other  hand,  birds  crush  the  fruits  and  discard  the  seed, 
which  is  surrounded  by  a  viscid  layer."  (Naturlichen  Pflan- 
zenfamilien.  Theil  III. )  Kerner  says:  "The  dissemination 
of  the  European  mistletoe  is  effected  as  in  all  Loranthaceas 
through  the  agency  of  birds,  thrushes  in  particular,  which  feed 
upon  the  berries  and  deposit  the  undigested  seeds  with  their  ex- 
crement upon  the  branches  of  trees."  (Nat.  Hist,  of  Plants, 
1 :  205,  1894.)  Keeble,  the  most  recent  observer  who  has  pub- 
lished upon  the  Loranthaceae,  says  :  "The  berry-like  fruits  of 
the  Loranths  are  technically  speaking  indehiscent;  yet  owing 
partly  to  growth  of  the  embryo,  partly  to  weakening  of  the 
fruit  wall  in  some  species,  this  latter  becomes  ruptured  on  the 
ripening  of  the  fruits,  e.  g.,  Loranthus  neelgherrensis  /,. 
cuneatus;  in  others  a  very  slight  pressure  is  sufficient  to  cause 
the  complete  extrusion  of  the  seed,  sometimes  basally,  some- 
times apically.  In  all  cases  the  seed  slips  out,  but  in  Viscum 
orientale  Willd.,  a  gentle  pressure  will  cause  the  fruit  wall 
to  crack  and  the  seed  to  be  jerked  out."  ( Observations  on  the 
Loranthaceae  of  Ceylon,  Trans.  Linn.  Soc.  Lond.  2nd  Ser. 
Bot.  5  :  Pt.  3,  p.  97,  1896.) 

In  view  of  the  above  statements  it  is  safe  to  conclude  that 
Razoumofskya  is  the  only  Loranth  furnished  with  a  mechanism 


MacDougal:    DISTRIBUTION  OF  KAZOUMOFSKYA  UOKUSTA.  171 

for  the  expulsion  of  the  seeds  from  the  berries  without  the  in- 
tervention or  cooperation  of  outside  factors.  The  expulsory 
mechanism  is  best  seen  in  a  longitudinal  section  of  the  berry. 
The  base  of  the  berry  is  joined  to  the  stalk  by  a  scission  layer 
several  cells  in  thickness.  The  outer  coat  is  firm  and  smooth, 
and  is  composed  of  an  epidermal  layer  with  the  outer  wall  ex- 
tremely heavy  and  cuticularized.  Beneath  the  epidermis  is  a 
mass  of  parenchymatous  tissues,  the  outer  portion  of  which  is 
slightly  palisaded  and  containing  chlorophyll,  the  inner  layer 
showing  only  starches  and  sugars.  Immediately  internal  is  the 
fibrovascular  framework  which  fuses  at  the  apex  in  a  solid  mass 
of  mechanical  tissue.  Lying  inside  the  fibrovascular  strands, 
and  continuous  with  the  parenchymatous  tissue  external  to  it  is 
also  a  mass  of  similar  thin-walled  elements  of  ovoid  or  cylindri- 
cal form  rich  in  carbohydrates.  These  cells  have  their  axes  at 
right  angles  to  the  surface  of  the  berry.  The  second  layer  in- 
ternal to  the  fibrovascular  tissue  is  the  expulsory  layer,  consist- 
ing of  very  long  thin-walled  cylindrical  tubes  with  their  axes 
parallel  to  the  long  axis  of  the  berry  at  the  apex  of  the  seed  or 
variously  inclined  from  this  position  according  to  the  location, 
but  all  so  arranged  that  their  longitudinal  expansion  would  tend 
to  force  the  seed  out  of  the  mouth  of  the  sac  formed  by  the 
berry.  Immediately  coating  the  seed  is  a  layer  of  globoid  cells 
with  thick  mucilaginous  contents.  The  seed  has  the  form  of  a 
modern  rifle  bullet,  conical  at  the  basal  end  and  truncate  at  the 
apical  end,  with  a  general  cylindrical  outline.  The  scission 
layer  appears  to  cut  into  the  mucilaginous  layer  or  at  least  very 
nearly  so  in  the  mature  berry.  During  the  ripening  period  the 
contents  of  the  expulsory  layer  undergo  such  chemical  changes 
as  to  give  the  contents  a  very  high  isotonic  coefficient.  The 
consequent  osmotic  attraction  of  water  into  this  layer  sets  up  a 
turgescence  which  could  not  be  measured,  but  which  probably 
amounted  to  many  atmospheres.  The  steady  increase  of  the 
turgidity  of  the  expulsory  layer  brings  the  tension  to  the  breaking 
strain  of  the  scission  layer,  and  its  sudden  and  complete  rupture 
permits  the  full  force  of  the  pressure  to  act  upon  the  seed,  send- 
ing it  to  a  distance  of  two  or  three  meters.  The  entire  arrange- 
ment is  that  of  a  mortar  cannon. 

The  muzzle  of  the  gun  is  sealed  by  the  stalk,  and  the  charge 
amounts  to  several  atmospheres,  which  is  allowed  to  act  upon 
the  seed  when  the  muzzle  is  freed.  The  firing  of  this  unique 


172  MINNESOTA    BOTANICAL    STUDIES. 

gun  may  result  from  the  overcoming  of  the  resistance  of  the 
restraining  layer  at  the  muzzle,  or  this  event  may  be  precipi- 
tated by  any  force  from  the  outside  which  would  result  in  the 
disturbance  of  the  scission  layer.  One  may  stand  under  a  pine 
tree  on  a  quiet  morning  and  hear  the  sharp  click  accompanying 
the  expulsion  of  the  seed  from  the  berries  at  irregular  intervals. 
If  the  branches  are  jarred  or  shaken,  however,  the  irregular  ex- 
plosions give  way  to  fusillades  by  which  nearly  all  of  the  berries 
on  a  plant  will  be  set  in  action  at  once. 

The  expulsion  of  the  seed  occurs  as  soon  as  the  berry  has 
broken  loose  from  the  stalk,  and  as  these  berries  were  originally 
in  all  positions  the  seeds  are  sent  out  in  all  directions.  The 
mucilage  adhering  to  the  seed  causes  its  attachment  to  the 
branches  or  other  bodies  it  may  strike.  In  this 
manner  dissemination  is  effected  throughout  a 

O 

cylindrical  space  about  seven  meters  in  diame- 
ter and    extending   downward   to   the   ground. 
The  only  localities  which  offer  suitable  condi- 
tions  for  the  germination  and    growth  of  the 
seeds,  however,  are  the  tips  of  branches  or  the 
shoots  of  young  trees  underneath.     It  is  to  be 
seen  that   no   animals  are  to  be  found  in  the 
habitat   of    the  parasite  which  would   in  ordi- 
nary usage  carry  the  seeds  to  these  locations. 
Seed  of  Razoumof-  The  only  part,  therefore,  that  animals  play  in 
skya  adhering  to  the    dissemination    of   the   seeds   would    be   in 
causing  the  discharge  of  the  berries,  a  matter 
of  no  direct  value,  since  they  are  capable  of  quite  as  efficient 
action  independently.     The  berry  of    Razoumofskya  is,  there- 
fore, to  be  classed  as   a   sling  fruit,  and  is  probably  the  only 
one  of  this  class  from  the  United  States  which  has  been  de- 
scribed, though  many  doubtless  exist. 

A  second  point  of  interest  in  this  plant  consists  of  a  fact  bear- 
ing upon  its  local  distribution.  During  the  course  of  some  re- 
cent field  work  in  northern  Arizona  the  writer  found  that 
Razoumofskya  was  most  successful  in  its  attacks  on  the  pine 
trees  along  the  rims  of  canons  or  along  the  brows  of  hills  or 
margins  of  mesas.  A  study  of  the  meteorological  conditions 
shows  that  this  method  of  distribution  has  a  direct  connection 
with  the  vertical  movements  of  the  air. 

As  the  air  resting  on  lowlands  in  canons  or  valleys  is  warmed 


MacDougal:   DISTRIBUTION  OF  RAZOUMOFSKYA  ROBUSTA.  173 

"by  radiation  during  the  time  of  exposure  to  the  sun's  rays,  it 
rises  and  expands.  During  the  ascent  some  heat  is  converted 
into  the  work  necessary  in  expansion,  causing  a  cooling  of  one 
degree  Fahrenheit  for  every  one  hundred  and  eighty-eight  feet 
of  elevation.  The  decrease  in  temperature  lowers  the  dew  point 
or  increases  the  relative  humidity,  a  matter  of  very  great  im- 
portance to  germinating  seeds  and  transpiring  leaves.  Razou- 
inofskya  is  especially  abundant,  precisely  at  the  places  where 
the  effect  of  the  ascending  humid  currents  of  air  is  greatest, 
along  the  margins  of  hills  and  mesas  and  the  rims  of  canons. 
This  is  very  noticeable  along  the  Grand  Canon  of  the  Colorado 
river,  in  the  Coconino  Forest  reserve,  where  the  air  rising 
more  than  a  vertical  kilometer  from  the  riverbed  pours  across  the 
pine-covered  mesa  at  a  much  lower  temperature  and  very  much 
nearer  the  dew  point  than  the  body  of  air  which  it  replaces. 
In  its  rise  it  has  lost  heat  at  the  normal  adiabatic  rate  to  the 
amount  of  about  twenty-five  degrees  F.,  and  has  undergone  a 
great  variation  with  respect  to  the  dew  point.  As  a  consequence 
of  the  increased  humidity  favorable  to  germination,  the  pines 
near  the  rim  of  the  canon  are  most  thickly  infested  with  the 
parasite  over  a  belt  one  to  four  or  five  kilometers  in  width  run- 
ning parallel  to  the  margin.  One  may  walk  through  the  forest 
and  note  the  decreasing  abundance  of  Razoumofskya  as  the  dis- 
tance from  the  canon  increases. 

In  recapitulation  of  the  facts  adduced  in  this  note  it  is  to  be 
said  that  the  berries  of  Rasoumofskya  are  to  be  classed  as  sling 
fruits,  the  only  one  from  North  America  hitherto  described,  and 
that  this  genus  is  the  only  one  of  the  Loranthaceas  furnished 
with  means  of  seed-dissemination  independent  of  gravity  and 
animals.  The  writer  also  believes  that  he  is  justified  in  an- 
nouncing the  discovery  of  the  influence  of  vertical  air-currents 
upon  the  distribution  of  plants,  and  that  this  factor  must  be  taken 
into  account  in  the  consideration  of  the  boundaries  of  zones  in 
mountainous  regions  or  those  with  irregular  topography. 

EXPLANATION  OF   PLATES. 

Plate  XV.  D.  Staminate  plants  of  Razoumofskya.  B.  Pistillate 
plants  with  mature  berries.  The  distortion  of  the  branch  of  the  host 
is  plainly  shown. 

Plate  XVI.  A.  Pinus  ponderosa  dying  from  the  effects  of  the  para- 
site, photograph  of  a  specimen  growing  on  the  extreme  edge  of  the  rim  of 
the  Grand  Canon  of  the  Colorado,  June,  1898.  C.  Specimen  of  Pinus 
ponderosa  showing  drooping  of  branches  attacked  by  Razoumofskya. 


VOL.   II. 


MINNESOTA    BOTANICAL    STUDIES. 


PART    II. 


PLATE    XV. 


VOL.    II. 


MINNESOTA    BOTANICAL    STUDIES. 


PART    II. 


D 


B 


PLATE    XVI. 


XIII.     OBSERVATIONS    ON   CONSTANTINEA. 


E.  M.  FREEMAN. 


History  and  Literature. — The  earliest  mention  of  the  red 
seaweed  now  classified  under  Constantinea  is  found  in  Gmelin's* 
Historia  Fucorum  published  in  1768,  in  which  he  describes 
Fucus  rosa-marina  from  the  material  collected  by  G.  W.  Steller 
during  the  years  1742-1745  at  Kamtschatka.  The  description 
is  as  follows:  "  Peculiare  sistet  haec  planta  fuci  specimen, 
cujus  exemplum  aliud  in  omni  reliqua  fucorum  historia  non  oc- 
currit.  Caulis  teres  est,  carnosus,  pennae  anserinas  crasitie, 
ramis  sibi  similibus,  quibus,  tanquam  totidem  pendiculis,  adpli- 
cantur  verticillatim  folia  petaloidea  terna  vel  plura,  rotunda, 
concava,  circulo  in  centre  notata,  pulchre  expansa,  plerumque 
sissa,  ramo  per  ilia  penetrante,  exeunte,  et  pollicis  dimidii  inter- 
vallo  nova  fronde  priori  simili,  prolifico,  tertia  nunnunquam 
pari  ratione  accedente.  Petala  convoluta  pulchre  repraesentant 
flores  polypetalos,  ut  Rosam,  anemonen,  cet.  Substantia  tota 
gelatinoso-membranacea,  aqua  dissoluenda,  pellucida.  Color  e 
rubro  flavescens.  Magnitude  semipedalis.  Locus.  Circa 
Lapatka  inter  spongias  ad  Kamtschatcam  occurrit." 

Such  terms  as  "  petala  convoluta,"  "  flores  polypetalos,"  etc., 
show  what  a  profound  impression  the  superficial  resemblance  of 
the  described  plant  to  a  rose  had  made  upon  the  author. 

In  the  years  1826-1829  the  Russian  vessel  Seniavin,  Fr. 
Liitke,  Captain  by  the  order  of  Czar  Nicolaus  I.,  sailed  through 
Russian  waters  and  collected  a  large  amount  of  valuable  algo- 
logical  material.  The  results  were  published  in  1840  by  Pos- 
tels  and  Ruprecht  in  their  Illustrationes  Algarum  in  itinere, 
etc.  The  authors  in  their  preface  to  this  work  state  that  the  ^ 
collections  of  H.  Mertens  and  the  plates  of  Alex.  Postels  form 
the  basis  of  the  entire  work.  The  genus  Constantinea  is  here 
described,  founded  upon  Gmelin's  Fucus  rosa-marina,  and  three 
species  are  recognized,  Constantinea  rosa-marina^  C.  sitchensis 
and  C.  reniformis. 

*Gmelin,  S.  G.     Hist.  Fuc.  102.  pl.j,f.2,  20.     1768. 


176  MINNESOTA    BOTANICAL    STUDIES. 

According  to  the  descriptions  the  first  two  species  of  Constan- 
tinea  differ  in  the  length  of  stipe  between  annuli,  the  mode  of 
branching  of  the  stipe  and  in  the  nature  of  the  edge  of  the  frond. 
The  following  are  also  noted :  Constantinea  rosa-marina  is  the 

O 

smaller  (one-half  foot  or  less) ;  branched  even  to  the  base  ;  termi- 
nal frond  two  inches  in  diameter,  laciniate  (lacinise  3-6),  rarely 
remaining  entire  ;  2—4  laminae  under  the  terminal  one  laciniate 
in  a  similar  manner.  C.  sitchensis  has  solitary  terminal  fronds 
at  the  apices  of  the  branches  ;  the  fronds  are  4-6  inches  in 
diameter,  entire  but  laciniate  when  older ;  young  fronds  are 
often  8  lines  in  diameter. 

In  these  supplementary  descriptions  all  differences  are  com- 
promised except  the  following:  length  of  the  "  internode  "  of 
the  stipe  ;  the  number  of  fronds  on  each  branch  ;  the  method  of 
branching  of  the  stipe  and  the  difference  in  size.  The  tetra- 
spores  of  C.  sitchensis  alone  are  described.  The  "  gongyli 
rotundi  "  as  Kiitzing*  has  since  pointed  out  are  but  ordinary 
cells  of  the  intermediate  layers  of  the  frond.  Postels  and  Ru- 
precht  also  mention  Constantinea  reniformis,  a  rare  Mediterran- 
ean plant  supposedly  of  this  genus. 

In  1843  Zanardinif  described  the  C.  reniformis  of  Postels 
and  Ruprecht  under  the  name  of  Neurocaulon  foliosuni  from 
material  collected  on  the  shores  of  Dalmatia  in  the  Adriatic.  In 
the  same  year  KiitzingJ  called  attention  to  the  incorrect  view  of 
Postels  and  Ruprecht  concerning  the  "  gongyli  rotundi  "  and  to 
the  great  similarity  in  vegetative  structure  but  great  difference 
in  outward  appearance  and  tetraspore  formation  between  Con- 
stantinea and  Euhymenia.  He  describes  the  tetraspores  and 
states  that  the  cystocarps  are  unknown.  His  statements  are 
evidently  based  solely  on  Postels,  and  Ruprecht's  observations. 

In  Species  Algarum,  1849, §  the  two  arctic  species  of  Postels 
and  Ruprecht  are  described  under  Neurocaulon  as  N.  rosa- 
marina  and  N.  sitchensis. 

Two  years  later  J.  Agardh  ||  accepts  the  Postels  and  Ruprecht 
generic  name  of  Constantinea  and  adds  to  the  previous  descrip- 
tion of  the  vegetative  parts  and  tetraspores  that  of  the  cysto- 


*  Kiitzing.     Phyc.  Gen.  400.     1843. 
tZanardini.     Saggio  class.  49.     1843. 
t  Kiitzing.     1.  c. 

§  Kiitzing.     Spec.  Algar,  744.      1849. 

||  Agardh,  J.     Spec.  Gen.  et  Ord.  Algar.  2  :  295.      1851. 


Freeman :   OBSERVATIONS  ON  CONSTANTINEA.  177 

carps.  Since  no  new  observations  nor  collections  of  the  arctic 
species  are  cited,  his  generic  description  of  the  cystocarp  is 
probably  based  upon  Constantinea  reniformis,  the  Mediter- 
ranean species.  The  cystocarps  are  described  as  "  kalidia  in 
media  fronde  numerosa,  clausa,  disruptione  partis  ambientis 
demum  liberata,  nucleolis  pluribus  composita ;  nucleoli  intra 
periderma  gelatinosum  hyalinum  gemmidia,  nullo  ordine  dis- 
posita  foventes."  The  zonate  division  of  the  tetraspores  is 
noted.  The  collections  of  C.  reniformis  cited  are  :  In  the  Ad- 
riatic sea  on  the  shores  of  Dalmatia  (Meneghini !  and  Zanar- 
dini!)  and  in  the  Mediterranean  sea  at  Cette  (Salzman  !)  and 
at  Marseilles  (Solier!).  Nothing  new  is  added  concerning  C. 
sitchensis  and  C.  rosa-marina,  but  C.  reniformis  is  fully  dis- 
cussed. The  latter  had  been  collected  also  by  Mertens  and  de- 
scribed by  him  under  the  name  of  Kalymenia  reniformis.  In 
1822  it  was  described  by  Agardh  *  under  the  name  of  Haly- 
mcnia  reniformis,  and  Postels  and  Ruprecht  f  describe  it  as  a 
third  species  of  Constantinea — C.  reniformis.  In  1822  J. 
Agardh  \  stated  that  the  tetraspores  of  C.  reniformis  had  not 
been  found. 

Harvey's  description  of  Constantinea  in  1858  §  is  based  upon 
the  observations  and  literature  cited  above.  The  similarity  in 
structure  and  the  difference  in  external  form  and  in  position  of 
tetraspores  is  noted.  The  branching  of  the  stipe  is  described 
as  at  first  irregular  but  later  dichotomous ;  the  dichotomy,  how- 
ever, is  often  lost  in  the  abortion  of  one  branch. 

In  1862  in  a  notice  of  a  collection  of  algae  made  by  Dr. 
David  Lyall  at  Vancouver  Island  in  the  years  1859-1861,  by 
W.  H.  Harvey  ||  specimens  of  Constantinea  sitchensis  with  torn 
laminse  which  were  probably  six  to  eight  inches  in  diameter 
when  perfect  were  reported  "  adrift  on  the  beach  at  Victoria 
harbor."  And  Harvey  observes  that  "perhaps  this  is  only  a 
luxuriant  state  of  Constantinea  rosa-marina." 

In  Kiitzing's  work  of  1867^  the  genus  is  again  described  un- 
der Neurocaulon  and  N.  foliosum  and  N.  rosa-marina  are 
mentioned.  The  work  of  Postels  and  Ruprecht  is  not  cited. 

*  Agardh.     Spec.  Alg.  201.     1822. 

t  Postels  and  Ruprecht.     111.  Alg.  17.     1840. 

%  Agardh,  J.     1.  c. 

§  Harvey,  W.  H.     Nereis  Boreali-Americana  2  :    173.     1853. 

[  Harvey,  W.  H.     Journ.  Proc.  Linn.  Soc.  Bot.  6 :  172.     1862. 

I  Kiitzing.     Tab.  Phyc.  17  :  24.  pi.  8$.     1867. 


178  MINNESOTA    BOTANICAL    STUDIES. 

Mention  is  again  made  of  the  genus  by  J.  Agardh  in  1876.* 
He  had  seen  specimens  of  C.  rosa-marina  from  the  Museum  of 
St.  Petersburg  and  also  a  plant  from  Californian  shores  which 
he  referred  to  C.  sitchensis;  but  on  account  of  inability  to 
satisfy  himself  as  to  the  structure  of  the  fruiting  bodies  he  based 
his  descriptions  upon  those  of  Postels  and  Ruprecht. 

In  1885  Constantinea  thiebauti  was  described  by  Bornetf  from 
a  single  specimen  collected  at  Majunga  on  the  north  coast  of 
Madagascar.  Bornet  very  properly  calls  attention  to  the  re- 
markable range  which 'this  genus,  with  the  addition  of  his  new 
species  would  enjoy :  C.  rosa-marina  and  C.  sitchensis  in  arctic 
seas,  C.  reniformis  in  the  deep  waters  of  the  Mediterranean  and 
C.  thiebauti  in  the  tropical  waters  of  the  Indian  ocean. 

In  1891  there  appeared  in  the  Botanical  Magazine  of  Tokyo 
"Remarks  on  some  algae  from  Hokkaido  "$  in  which  Con- 
stantinea  sitchensis  Post,  and  Rupr.  (?)  is  mentioned,  but  the 
position  of  the  described  plants  in  the  genus  Constantinea  is  ad- 
mitted to  be  doubtful. 

In  "Die  Natiirlichen  Pflanzenf  amilien  "  §  the  genus  under- 
goes rearrangement.  C.  reniformis  is  restored  to  the  genus 
Neurocaulon  of  Zanardini,  where  probably  should  also  be  placed 
Bornet's  C.  thiebauti.  The  reproductive  bodies,  as  well  as  the 
vegetative  structure  of  C.  rosa-marina  and  JV.  foliosum,  had 
been  studied  personally  by  Schmitz.||  The  classification  of 
Schmitz  and  Hauptfleisch  includes  under  Constantinea  the  species 
C.  sitchensis  and  C.  rosa-marina  and  under  Netirocaulon  the 
single  species  N.foliosum.  I  have  no  certain  knowledge  as  to 
whether  Schmitz  had  or  had  not  in  possession  any  material  of 
C.  sitchensis. 

Collection. — In  August,  1897,  and  again  in  the  summer  of 
1898  collections  of  a  species  of  Constantinea,  reported  as  C. 
sitchensis^  were  made  by  Miss  Josephine  E.  Tilden,  and  it  is 
upon  this  material  that  the  following  observations  are  based  : 

On  August  3,  1897,  specimens  of   C.  sitchensis  were  found 

*  Agardh,  J.     Spec.  Alg.  3  :  225.     1876. 

t  Bornet.     Alg.  de  Mad.  Bull.  Soc.  Bot.  de  France  32  :  18.  /  /.  2.     1885. 

JOkamura,  R.  Remarks  on  some  algae  fron  Hokkaido.  Bot.  Mag.  Tokyo. 
5=  333-336-  1891. 

§  Schmitz  and  Hauptfleisch.  Nat.  Pflanz.  I.  Teil.  Abt.  2:  517,  519,  520,  525. 
1897. 

II  Schmitz.  Sys.  Ubers.  der  bisher  bekannten  Gattungen  der  Florideen.  Flora 
72  :  436.  1889. 


Freeman :   OBSERVATIONS  ON  CONSTANTINEA.  179 

growing  upon  holdfasts  of  Nereocystis  lutkeana  in  8  fathoms 
of  water  at  Channel  Rocks,  near  Seattle,  Washington.  Dur- 
ing the  summer  of  1898  large  collections  were  made  at  many 
of  the  stations  where  Dr.  Lyall  collected  in  1859-1861.  Col- 
lections were  made  at  the  following  places:  (i)  Fairhaven, 
Washington.  May  25.  Washed  up  on  the  beach.  (2)  Near 
Minnesota  reef,  San  Juan  island,  Washington.  June  5.  At- 
tached to  stones  on  a  flat,  sandy  beach.  This  and  the  three 
following  were  found  just  below  lowest  tide.  (3)  Near  Friday 
Harbor,  San  Juan  island,  Washington.  June  5.  Attached  to 
rocks  on  rocky,  steep  beach.  (4)  Oak  Bay  (a  suburb  of  Vic- 
toria), British  Columbia.  July  i.  On  a  sandy  beach.  (5) 
Esquimalt,  British  Columbia.  July  2.  Attached  to  rocks. 
The  first  and  last  two  collections  contained  abundant  tetraspore 
material. 

Preservation. — The  material  collected  in  1897  was  killed  and 
preserved  in  80%  alcohol.  Owing  to  the  small  amount  of  this 
material  and  to  the  better  condition  of  that  collected  in  1898, 
all  of  the  following  drawings  except  Fig.  i  have  been  made 
from  the  1898  material.  The  larger  part  of  it  was  killed  and 
preserved  in  a  2  per  cent,  formalin  solution  in  sea  water.  In 
this  the  color  was  very  well  preserved.  The  firmness  of  the 
tissues,  however,  suffered  considerably  more  in  the  formalin- 
solution  material  than  did  that  of  the  alcohol  material  of  1897. 
The  formalin  material  was  still  sufficiently  firm  to  admit  of  very 
satisfactory  manipulation.  Still  other  plants  were  preserved  in 
camphor  water,  and  some  in  i  per  cent,  chromic  acid  solution. 
The  camphor  material  lost  its  color  almost  as  completely  as  the 
alcoholic.  It  preserved,  however,  a  great  firmness,  which  ren- 
dered the  tissues  excellent  for  section  cutting,  and  especially 
for  hand  sections.  The  gelatinous  cell  walls,  however,  were 
so  cleared  that  they  were  not  as  easily  defined  as  in  the  formalin 
material.  The  chromic  acid  collections  were  in  a  poor  state  of 
preservation ;  the  tissues  were  very  soft,  the  cell  walls  almost 
invisible  and  the  contents  usually,  at  least  partially,  disorganized. 

Methods. — Various  methods  were  employed  in  cutting  the  tis- 
sues. A  part  of  the  material  was  transferred  directly  from 
water  to  20  per  cent,  glycerine,  thence  to  a  gum  arabic  solution 
upon  an  Osterhout  freezing  chamber.*  Material  was  some- 
times placed  directly  from  the  sea  water  on  the  freezing  chamber 

*  Osterhout,  W.  J.  V.    Bot.  Gaz.  21  :  195.     1896. 


180  MINNESOTA    BOTANICAL    STUDIES. 

with  results  almost,  if  not  equal,  to  those  obtained  when  it  was 
passed  first  into  20  per  cent,  glycerine.  When  the  sections  were 
removed  from  the  knife  they  were  placed  in  20  per  cent,  glycer- 
ine, and  from  this  into  50  per  cent,  and  then  into  absolute  glycer- 
ine ;  or  from  20  per  cent,  glycerine  to  water  and  then  through  the 
alcohols  to  an  alcohol  stain  or  directly  into  a  water  stain.  The 
freezing  device  mentioned  above  proved  very  satisfactory  in 
many  respects.  The  tissues  can  be  frozen  in  a  minute's  time, 
are  held  firmly  in  place,  and  the  gum  arabic,  when  of  the  proper 
consistency,  is  an  excellent  imbedding  medium.  The  difficulty 
experienced  with  this  method  of  cutting  such  delicate  tissues  as 
are  found  in  many  of  the  seaweeds  lies  in  the  handling  of  sec- 
tions after  they  are  cut.  Especially  is  this  true  when  it  is  desir- 
able to  stain  the  sections  and  when  they  must  be  transferred 
through  several  per  cents,  of  alcohol  and  glycerine.  With  such 
tissues  as  are  found  in  the  frond  of  Constantinea^  where  they 
part  with  great  ease,  the  difficulty  is  augmented,  and  it  was 
found  almost  impossible  to  preserve  thin  sections  whole.  An 
attempt  was  made  to  obviate  this  difficulty  by  mounting  the  sec- 
tions directly  from  the  microtome  knife  into  glycerine  jelly  at  a 
temperature  sufficient  to  keep  the  jelly  semi-fluid.  This  was  in 
part  an  improvement,  but  necessitated  the  mounting  of  many 
worthless  sections.  When  sections  unstained  were  mounted  in 
glycerine  or  glycerine  jelly,  the  great  transparency  of  the 
swollen  cell  walls  added  a  new  difficulty. 

Material  was  also  passed  into  paraffin  and  stained  on  the  slide 
and  mounted  in  Canada  balsam  by  the  usual  methods.  Ex- 
treme care  was  found  necessary,  on  account  of  the  delicacy  of 
the  tissues,  to  make  the  stages  from  one  fluid  to  another  by  very 
gradual  changes.  I  was  unable  to  prevent  a  partial  shrinkage 
of  the  cell  contents.  Sections  by  this  method  were  otherwise 
quite  satisfactory,  having  the  advantage  of  use  in  serial  work. 
Staining  is  necessary  in  this  method  since  the  sections  become 
almost  invisible  in  Canada  balsam. 

Sections  cut  freehand  with  a  razor  or  in  a  hand  microtome 
with  the  material  imbedded  in  pith  have  furnished  most  of  the 
material  from  which  the  accompanying  plates  are  drawn.  Suf- 
ficiently thin  sections  were  obtainable  in  this  way  with  the  great 
advantage  of  certainty  as  to  the  normal  condition  of  the  tissues 
and  of  speed  in  preparation.  A  large  number  of  such  sections 
can  be  cut  and  preserved  in  2  per  cent,  formalin  for  a  long  time 
and  are  ready  for  use  at  any  moment. 


Freeman:   OBSERVATIONS  ON  CONSTANTINEA.  181 

A  number  of  staining  fluids  were  used,  section  staining, 
either  on  or  off  the  slide  proving  more  satisfactory  than  stain- 
ing in  toto.  Aniline  stains  were  used  almost  exclusively.  The 
following  were  employed  with  at  least  some  degree  of  success  : 

Aniline  blue  :  Stains  the  gelatinous  wall  of  the  cells  pale 
blue,  the  chromatophores  of  the  cortical  cells  deep  blue  and  the 
contents  of  the  tetraspores  and  of  the  paraphyses  light  blue. 
The  central  part  of  the  pyrenoids  of  the  endophytic  alga  which 
is  usually  present  in  these  collections  takes  on  a  light  blue.  The 
best  results  were  obtained  with  a  40  or  60  per  cent,  alcoholic 
solution,  acting  from  24  to  36  hours.  For  sections  on  the  slide 
5  to  10  minutes  in  a  strong  60  per  cent,  alcoholic  solution  was 
usually  sufficient.  My  best  staining  results  were  obtained  from 
aniline  blue. 

Methylen  blue  :  Stains  the  cell  walls,  especially  the  outer 
portions,  which  become  quite  clearly  defined.  Sections  were 
stained  in  a  strong  95  per  cent,  alcoholic  solution  from  5  to  10 
minutes. 

Fuchsin  :  Best  results  were  obtained  from  sections  left  in  a 
10  per  cent,  alcoholic  solution  36  hours.  The  gelatinous  walls 
were  stained  light  red  while  the  protoplasm  of  the  paraphyses 
and  chromatophores  of  the  cortical  cells  took  up  a  deep  carmine 
red. 

Delafield's  hasmatoxylin  :  Same  strength  and  time  as  fuchsin. 
The  granular  contents  of  the  cells  are  stained  a  reddish  purple. 

Several  other  stains  were  used,  but  without  success.  In  a 
weak  solution  of  iodine  in  potassium  iodide  the  granules  of  the 
cells  of  the  middle  and  intermediate  layers  assume  at  first  a  yel- 
lowish-brown tinge  which  finally  deepens  to  an  amethyst  purple. 
Stained  in  a  strong  solution  for  a  few  moments  these  areas  take 
on  a  dense  violet  color. 

Gross  Anatomy. — In  general.form  Constantinea  presents  sev- 
eral interesting  peculiarities.  (Figs.  I— 6.)  The  plant  is  differen- 
tiated into  a  cylindrical  stipe  and  peltate  frond  at  its  summit.  It 
is  of  a  purple-reddish  color,  stands  upright  in  the  water  below 
lowest  low  tide  mark,  and  the  texture  is  quite  firm  and  brittle. 
It  is  somewhat  gregarious  in  habit.  The  stipe  is  "  terete, 
branched,  ringed  and  the  apex  of  each  branch  expanded  into  an 
orbicular  peltate  lamina;  stipe  1-4  mm.  in  diameter,  2-8  cm. 
in  length  ;  lamina  2  cm. -3  dcm.  in  diameter."  (Tilden,  Am. 
Alg.  no.  203.  1897-)  The  laminae  have  numerous  minute 


182  MINNESOTA    BOTANICAL    STUDIES. 

dark  brownish  spots  on  their  upper  surfaces.  The  older  fronds 
are  irregularly  perforated  with  circular  holes  and  are  more  or 
less  torn  at  the  edges  (Fig.  3).  When  young  the  fronds  are  en- 
tire (Fig.  2).  As  the  plant  continues  its  growth  the  stem  pushes 
up  through  the  center  of  the  frond,  forming  a  new  growing 
point  which  elongates  into  an  internode  and  soon  forms  another 
frond  at  its  summit.  The  old  frond  then  falls  off,  leaving  an 
annulate  marking  on  the  stipe  (Figs.  5,6).  There  is  often,  espe- 
cially in  the  youngest  portions  of  the  oldest  plants,  a  formation  of 
two  growing  points  at  the  center  of  each  lamina  giving  rise  to 
a  dichotomy  of  the  stipe  (Fig.  5).  Irregularity  in  the  develop- 
ment of  these  growing  points  gives  rise  to  an  irregular  system 
of  branching  (Fig.  5).  Growing  points  are  not  confined  to  the 
bases  of  laminae  but  may  be  found  occasionally  arising  from  other 
portions  of  the  stipe.  There  are  often  a  group  of  small  branches 
formed  upon  the  holdfast  (Fig.  i).  The  latter  is  a  disc-like  body 
arising  as  an  expanded  portion  of  the  stipe  at  its  base  and  usu- 
ally concave  below  (Fig.  22). 

Upon  the  lower  surface  of  the  fronds  especially  the  larger 
ones,  can  be  seen  distinct  radial  striations  caused  by  small  ridges 
running  from  the  region  near  the  stipe  toward  the  edge  of  the 
frond  (Fig.  4).  These  mark  the  course  of  bundles  of  elongated 
narrow  filaments  (see  below).  The  nemathecia  which  bear  the 
tetraspores  are  found  only  on  the  lower  surface  of  the  frond. 
They  occur  more  abundantly  on  the  larger  fronds  and  are  often 
so  numerous  as  to  almost  completely  cover  the  under  surface. 
They  form  small  wart-like  bodies  of  whitish  color  and  gelatin- 
ous consistency. 

Minute  Anatomy. — As  is  well  known  the  thallus  of  the  red 
seaweed  is  a  group  of  dichotomously  branching  filaments  whose 
fused  branches  form  tissue-like  areas.  In  Constantinea  this 
method  of  branching  can  readily  be  seen  in  the  frond  but  par- 
ticularly so  in  the  cortical  area  of  the  stipe  (Fig.  18). 

Frond. — A  cross  section  of  the  frond  shows  three  areas  of 
cells  :  (a)  a  central  layer  of  loosely  woven  filaments  ;  (b)  an  in- 
termediate layer  of  large  approximately  spherical  cells  stuffed 
with  starch  granules  (Fig.  7) ;  (c]  a  corticcal  layer  of  pseudo-pa- 
renchymatous  cells. 

a.  There  are  in  the  central  area  filaments  of  large  cells, 
usually  slightly  elongated,  often  stretching  across  the  frond  and 
perpendicular  to  the  frond  surface  (Fig.  n).  These  cells  ap- 


Freeman:   OBSERVATIONS  ox  COXSTANTINEA.  183 

proach  the  large  cells  of  the  intermediate  layer  in  form  and  size, 
transitional  stages  between  them  being  abundant  (Figs.  8,  9,  10). 
Their  average  size  is  55  mic.  x  12  mic.,  but  they  may  attain 
75  x  16  mic.  A  cross  section  of  the  lower  part  of  the  frond, 
tangential  to  its  orbicular  outline,  shows  in  the  central  area  at 
more  or  less  regular  intervals  corresponding  to  the  external  stri- 
ation  on  the  lower  surface  of  the  frond,  a  number  of  bundles  of 
greatly  elongated  cells,  woven  into  strengthening  bundles  (Figs. 
7,  8,  9).  In  the  lower  region  the  general  course  of  these 
bundles  is  radial  in  the  frond.  Divergence  from  this  course  is 
found  in  the  upper  part  so  that  a  tangential  cross  section  of  the 
frond  shows  some  of  these  bundles  in  longitudinal  or  oblique 
section  (Fig.  9).  These  filaments  are  articulated;  the  cells  of- 
ten attain  a  length  of  325  mic.  They  average  about  8  mic.  in 
breadth.  Thin  sections  of  the  frond  often  contain  clean-cut  cir- 
cular spaces  where  the  bundles  of  elongated  cells  have  been 
pulled  out,  showing  the  compactness  of  the  bundles  and  indicat- 
ing for  it  a  strengthening  function.  The  bundles  vary  in  size  ; 
the  striations  on  the  lower  surface  of  the  frond  mark  only  the 
largest  bundles. 

b.  The  typical  cells  of  the  intermediate   area   are   spherical 
and  average  46  mic.  in  diameter.     They  are  packed  with  Flor- 
idian    starch  granules  which  turn  brownish  with  a  weak  KI  so- 
lution of  iodine  and  finally  purple  or  violet  in  a  strong  solution. 
Those  cells  toward  the  surface  of  the  frond  contain  ordinarily  a 
few  chromatophores,  usually  in  the  end  near  the  surface.     The 
cells  of  the  intermediate  layer  shade  off  towards  the  surface  into 
the  cortical  cells. 

c.  The  cortical  cells  are  characterized  by  a  compact  pseudo- 
parenchymatous  grouping   in  which  the  long  diameters   of  the 
but  slightly  elongated  cells  are  perpendicular  to  the  frond  sur- 
face.    Many  of  the  cells  are  approximately  cylindrical  or  pris- 
matic.    The  external  layer  is  of  a  sufficient  regularity  in  struc- 
ture and  form  to  recall  forcibly  an   epidermal  layer.     In  these 
the  outer  wall  is  rounded.     The  cells  of  the  cortical  layer  con- 
tain but  a  small  amount  of  starch,  and  this  is  found  in  the  cells 
adjoining  the   intermediate  layer.     Chromatophores  are,  how- 
ever, abundant  and  more  numerous  toward  the  frond  surface. 
Almost  all  of  them  are  to  be  found  in  the  first  three  or  four 
rows  of  cells  from  the  surface  (Fig.  10),  but  some  are  found  in 
still   deeper  layers.      They  occupy,  in  the  great    majority  of 


184  MINNESOTA    BOTANICAL    STUDIES. 

cases,  a  position  in  the  peripheral  end  of  the  cell.  Under  the 
microscope  they  appear  finely  rose-colored  pink  with  perhaps  a 
purple  tinge.  They  are  irregular  in  outline,  apparently  in  most 
cases  assuming  such  a  shape  as  will  allow  them  to  occupy  the 
peripheral  end  of  the  cell  to  best  advantage.  The  epidermal 
cells  average  about  13x5  mic.  and  the  chromatophores  about 
10  x  2  mic. 

Stipe. — (Figs.  14-21).  A  cross  section  of  the  stipe  shows  a 
similarity  in  structure  to  that  of  the  frond.  The  same  areas  are 
present  with  modifications,  however  (Fig.  14).  The  cortical 
area  is  composed  of  cells  more  elongated  than  those  in  the 
frond  and  is  a  larger  number  of  cells  deep.  Internally  these 
pass  into  the  large  cells  of  the  intermediate  area  (Fig.  19). 
Here  there  is  a  noticeable  difference  from  the  condition  in  the 
frond.  A  large  number  of  the  elongated  central  cells  find  their 
way  into  this  area  and  a  cross  section  of  the  stipe  shows  them 
in  cross,  oblique  and  even  longitudinal  section  between  the 
large  starch-containing  cells  of  the  intermediate  area  which 
stand  out  very  clearly  in  large  radial  filaments  (Fig.  21).  The 
central  area  in  the  stipe  is  a  very  compact  area  and  is  made  up 
of  a  large  number  of  thin  filaments  interwoven  in  a  very  com- 
plicated manner  (Figs.  20,  21).  These  cells  correspond  to  those 
of  the  elongated  filament  bundles  in  the  frond.  There  are  in 
the  stipe,  moreover,  a  small  number  of  larger  cells  also  elon- 
gated and  corresponding  to  the  cells  of  similar  shape  and  posi- 
tion in  the  frond.  In  cross  section  the  cut-off  ends  predominate 
(Fig.  20),  while  in  longitudinal  section  the  cut-off  ends  are  few  ; 
the  longitudinal  view  of  the  filaments  is  the  predominant  one 
and  the  intricate  weaving  (Fig.  21)  is  very  plainly  seen.  A 
longitudinal  section  through  the  annulate  portion  shows  the  ab- 
sence of  the  cortical  and  intermediate  layers  in  the  region  of 
the  annulus  indicating  the  continuity  of  these  areas  in  the  frond 
and  stipe  (Fig.  15). 

A  longitudinal  section  through  the  growing  point  shows  but 
two  areas  in  the  growing  region.  There  is  no  distinct  interme- 
diate area  although  a  number  of  large  cells  may  be  present 
(Fig.  1 6).  The  two  areas  are  the  cortical  in  which  the  fila- 
ments are  all  parallel,  perpendicular  to  the  stipe  surface  and 
pseudc-parenchymatous  in  character,  and  the  central,  which  is 
as  before  a  mass  of  densely  woven  elongated  filaments.  The 
end  of  the  intermediate  area  of  the  frond  can  readily  be  seen 
in  such  a  section  (Fig.  16). 


FrecDiau  :    <>I:M-:I;  VATIOXS  ox  CONSTANTINEA.  185 

Holdfast. — The  holdfast,  i.  e.,  the  expanded  portion  of  the 
stipe  at  its  base,  presents  a  similar  anatomical  structure  to  that 
of  the  stipe  with  the  exception  of  one  modification.  The  lower 
cortical  and  intermediate  areas  which  are  in  contact  with  the 
surface  to  which  the  plant  is  moored  are  changed  into  a  yellow- 
ish brown  disorganized  mass  which  probably  serves  as  a  cement- 
ing substance  in  attaching  the  plant  to  the  rocks.  Indications 
of  the  former  cell  structure  can  be  seen  in  occasional  cavities 
and  in  the  arrangement  of  these  cavities.  The  area  adjoining 
the  cement  layer  is  composed  largely  of  elongated  filaments 
(Fig.  24). 

Protoplasmic  Connections. — In  freshly  cut  material  the  proto- 
plasmic connections  between  cells,  is  very  plainly  to  be  seen  in 
many  of  the  starch  cells  of  the  intermediate  area.  They  are 
also  easily  seen  in  the  larger  cells  of  the  central  area,  as  well  as 
in  the  small  cells  of  the  same  area.  The  cortical  area  of  the 
stipe  furnishes  particularly  good  views  of  this  continuity  (Figs. 
i3,  18). 

Reproductive  Tract  (Figs,  u,  12). — I  have  been  unable  to 
find,  in  the  material  at  hand,  and  it  is  considerable  in  amount, 
any  trace  of  cystocarp  development.  The  following  description 
of  the  occurrence  and  structure  of  the  cystocarp  of  the  genus 
is  translated  from  Schmitz  and  Hauptfleisch,*  the  former  of 
whom  has  made  a  personal  study  of  the  cystocarp  of  C.  rosa- 
marina  (Schmitz,  1.  c.)  :  The  carpogonial  branches  and  aux- 
iliary cell  branches  are  distributed  in  the  fertile  portions  of 
the  frond  in  large  numbers  in  the  loosened  inner  portion  of  the 
inner  cortex  of  the  upper  side  of  the  leaf,  together  with  numer- 
ous vermiform  sterile  cellular  threads.  Cystocarps  distributed 
in  large  numbers  on  the  fertile  fronds,  comprising  a  broad  zone 
along  the  edge  of  the  frond  on  its  upper  surface,  imbedded  in 
the  much  loosened  inner  cortex  of  the  upper  surface  of  the 
frond,  swelling  out  into  an  arch  the  superposed  outer  cortex 
which  is  punctured  by  pores.  The  nucleus  of  the  form  of  a 
mulberry  and  pierced  by  single  strands  of  sterile  tissue.  Goni- 
molobes  separated  only  at  first,  later  confluent. 

Kutzing  f  states  that  the  "  gongyli  rotundi "  described  by 
Postels  and  Ruprecht  as  two  kinds  of  fruiting  bodies  in  Constan- 
tinea  rosa-marina  are  ordinary  cells  of  the  cubcortical  layer. 

*  Schmitz  and  Hauptfleisch.  In  Engl.  and  Prantl.  Nat.  Pflanz.  I.  Teil.  Abth. 
2.  520,  521.  1897. 

t  Phyc.  Gen.  400.     1843. 


186  MINNESOTA    BOTANICAL    STUDIES. 

Tetraspores  are  known  only  in  C.  sitchensis.  They  are  ob- 
long and  zonate,  lodged  in  nemathecia.  The  nemathecia  are 
found  exclusively  on  the  lower  surface  of  the  frond  more  abun- 
dant near  the  outer  half.  They  are  in  the  form  of  delicate, 
slightly-raised  "  wart-like,"  often  confluent  bodies  of  a  whitish 
color.  They  average  3.5-4  mm.  in  diameter  and  often  become 
so  numerous  that  they  completely  cover  a  very  large  part  of  the 
lower  surface  of  the  frond.  The  nemathecia  are  covered  by  the 
gelatinous  layer  on  the  surface  of  the  paraphyses.  The  para- 
physes  are  elongated,  narrow,  peripheral  cells  (Figs,  n,  12). 
The  tetraspores  arise  as  club-shaped  elongations  of  peripheral 
cells  between  the  paraphyses  and  are  divided  zonately  into  four 
chambers.  The  tetraspores  are  numerous  in  each  nemathecium. 
Their  average  size  is  108x22  mic. 

Endophyte. — In  a  large  majority  of  sections  and  upon  all 
material  examined,  are  present  peculiar  green  approximately 
spherical  bodies  imbedded  in  the  cortical  tissue  of  the  frond. 
They  are  endophytic  algae  and  probably  the  Chlorochytriiim  in- 
clusum  of  Kjellman.  In  general  they  are  pear-shaped  with 
their  small  end  toward  the  surface  and  the  cell  wall  at  that  end 
thickened.  They  are  greenish  in  color  and  contain  a  number 
of  conspicuous  pyrenoids,  the  central  areas  of  which  stain  very 
readily,  having  a  particular  affinity  for  aniline  blue.  The  proto- 
plasm is  denser  toward  the  small  end  where  the  cell  wall  is  also 
thick.  It  is  with  this  end  that  the  endophyte  breaks  through 
the  cortical  tissues  of  the  nurse  plant.  I  have  as  yet  been  un- 
able to  detect  any  zoospore  formation.  This  interesting  little 
endophyte  will  receive  a  more  complete  discussion  in  a  subse- 
quent paper. 

Conclusions. — The  material  upon  which  these  observations 
are  based  was  distributed  by  Miss  Tilden  as  Constantinea 
sitchensis  Post,  and  Ruprecht.  A  careful  comparison  of  it 
with  the  plates  and  descriptions  of  Postels  and  Ruprecht  shows 
however  that  the  plant  under  observation  might  as  well  perhaps 
be  placed  under  Constantinea  rosa-marina.  The  material 
agrees  in  almost  every  particular  with  C.  rosa-marina  hav- 
ing, however,  single  terminal  fronds  and  an  occasional  evi- 
dent dichotomy  of  branching.  The  differences  enumerated 
above  in  the  descriptions  of  the  two  species  can  hardly  be  con- 
sidered of  specific  importance.  The  length  of  the  internodes 
may  vary  considerably.  The  greater  part  of  the  material  under 


Freeman  :   OBSERVATIONS  ON  CONST ANTINEA.  187 

observation  contained  long  internodes.  This  material  was  col- 
lected in  late  summer.  The  number  of  fronds  on  a  branch  is 
also  given  specific  value  by  Postels  and  Ruprecht.  It  is  prob- 
ably of  important  significance  that  no  tetraspores  were  found 
by  these  authors  upon  C.  rosa-marina  but  that  large  numbers 
were  found  upon  C.  sitchensis.  Figure  6  represents  a  young 
frond  having  no  tetraspores  but  with  a  succession  of  fronds 
similar  lo  those  of  C.  rosa-marina,  while  almost  all  of  the  re- 
maining material  had  solitary  terminal  fronds  crowded  with 
tetraspores.  The  material  collected  in  May,  1897,  contains 
tetraspores. 

The  dichotomy  of  the  branching  of  C.  sitchensis  is  a  com- 
paratively late  development  in  the  growth  of  the  stipe  and  is 
not  seen  in  the  older  parts.  C.  sitchensis  is  further  described 
as  larger  in  all  parts  than  C.  rosa-marina,  though  actual  meas- 
urements given  do  not  accord  with  this.  These  facts  suggest  the 
probability  that  the  C.  sitchensis  of  Postels  and  Ruprecht  is  the 
late  summer  stage  of  C.  rosa-marina.  This  supposition  ex- 
plains satisfactorily  the  absence  of  tetraspores  in  the  one  and  of 
dichotomy  of  the  stipe  in  the  other,  the  comparative  lengths  of 
"  internodes"  and  the  difference  in  sizes  of  the  two  plants. 

The  observations  and  impressions  of  subsequent  writers  add 
additional  weight  to  this  view.  In  recounting  the  founding  of 
the  genus  by  Postels  and  Ruprecht  upon  the  Fucns  rosa-marina 
of  Gmelin,  Agardh  (1.  c.  1851)  observes  of  C.  sitchensis  "  novo 
consimili  adjecta  specie  "  (p.  294)  and  of  C.  rosa-marina  "  pras- 
cedente  (sitchensis)  videtur  proxima,  cum  nulla  alia  confun- 
denda"  (p.  296). 

Of  the  material  collected  by  David  Lyall  at  Vancouver  island 
"adrift  on  the  beach  at  Victoria  harbor"  and  reported  as  C. 
sitchensis,  although  corresponding  in  size  to  C.  sitchensis  Har- 
vey (1.  c.  1862)  remarks  "  perhaps  this  is  only  a  luxuriant  state 
of  C.  rosa-marina" 

From  these  facts  it  would  seem,  therefore,  highly  probable 
that  C.  sitchensis  and  C.  rosa-marina  are  but  different  forms  of 
the  same  plant,  and  since  the  work  of  Schmitz  has  removed  C. 
reniformis  to  the  genus  Neurocaulon  where  also  it  is  probable 
that  C.  thiebauti  should  be  classified,  that  Constantinea  is  a 
monotypic  genus,  with  Constantinea  rosa-marina  as  the  Only 
species. 


188  MINNESOTA  BOTANICAL  STUDIES. 

DESCRIPTION  OF  PLATES. 
Fig.  i.   Young  plant  with  group  of  young  branches  on  the  holdfast. 

Fig.  2.  Portion  of  the  plant  showing  the  entire  edge  and  form  of 
young  fronds  (from  dried  material),  x  y2. 

Fig.  3.  An  old  frond  showing  lacerated  border  and  perforations 
(from  dried  material),  x  %% 

Fig.  4.  A  frond  showing  striations  on  the  lower  surface  (dried 
specimen),  x^. 

Fig.  5.  Plant  with  fronds  almost  entirely  cut  away  showing  the 
growing  points,  the  annulations  of  the  stipe  and  the  dichotomous 
branching,  a.  annulations.  g-p>  growing  point,  f.  frond  cut  off. 

Fig.  6.   Small  branch  showing  a  rapid  succession  of  laminae,     x  J^  . 

Fig.  7.  Diagram  of  a  tangential  cross  section  of  the  frond.  c. 
cortical  area.  int.  intermediate.  e.  bundles  of  enlarged  filaments. 
/.  loosely  woven  cells  of  central  area. 

Fig.  8.  Cross  section  of  a  frond  showing  bundle  of  elongated  cells 
in  central  area  in  transverse  section.  Letters  as  above.  Drawn  with 
camera  lucida.  x  250. 

Fig.  9.  Cross  section  of  a  frond  showing  a  longitudinal  section  of 
a  part  of  a  bundle  of  elongated  central  cells.  Drawn  with  camera 
lucida.  x  250. 

Fig.  10.  Cross  section  of  a  frond  stained  for  a  minute  in  a  strong 
solution  of  I  in  KI.  ch.  chromatophores.  st.  starch  grains.  Drawn 
with  camera  lucida.  X345- 

Fig.  ii.  Cross  section  of  a  frond  through  a  nemathecium.  cent. 
central  layer,  p.  paraphyses.  /.  tetraspores.  Cells  are  drawn  only 
in  outline.  Contents  have  been  omitted.  Camera  lucida.  xS3- 

Fig.  12.  Cross  section  of  a  frond  through  a  nemathecium  showing 
tetraspores.  Drawn  from  a  glycerine  mount  in  which  the  gelatinous 
walls  became  almost  entirely  obliterated.  The  walls  are,  therefore, 
omitted  except  around  the  tetraspores.  x  250. 

•Fig-  13-  Cells  from  central  area  showing  protoplasmic  connec- 
tions. Camera  lucida.  x  250. 

Fig.  14.  Diagram  of  cross  section  of  stipe  in  internode.  Letters  as 
in  frond. 

Fig.  15.   Diagram  of  longitudinal  section  of  a  stipe  through  a  node. 

Fig.  1 6.  Diagram  of  a  longitudinal  section  of  the  stipe  through  a 
growing  point. 

Fig.  17.  Peripheral  cells  from  cross  section  of  stipe  showing  the 
striations  in  the  outer  gelatinous  covering,  x  345. 

Fig.  1 8.   Filament  from  a  cross  section  of  the  stipe  in  the  cortical 


/•"rcenian :   OBSERVATIONS  ox  CONSTANTINEA.  189 

area  showing  the  dichotomy  of  the  branching.  Camera  lucida  for  an 
outline,  x  250. 

Fig.  19.  Cross  section  of  stipe  including  the  cortical  and  the  begin- 
ning of  intermediate  layer.  The  cut  off  ends  of  elongated  central  fila- 
ments are  seen  in  the  intermediate  area.  Camera  lucida.  x  130. 

Fig.  20.  Cross  section  of  stipe  in  the  central  area ;  shows  the  pre- 
dominance of  elongated  filaments,  x  250.  Outlined  with  camera 
lucida. 

Fig.  21.  Longitudinal  section  of  a  stipe  at  the  inner  edge  of  the 
cortex  ;  shows  the  complication  of  the  elongated  filaments.  Outlined 
with  camera  lucida.  x  250. 

Fig.  22.  External  view  of  holdfast  seen  from  below,     x  */£ . 

Fig.  23.  Diagram  of  cross  section  of  a  holdfast,  m.  layer  of  disor- 
ganized mass  on  lower  surface  by  which  the  holdfast  adheres. 

Fig.  24.  Lower  portion  of  the  cross  section  of  the  holdfast  showing 
the  attaching  layer  m.  Camera  lucida.  x  250. 

Fig.  25.  Diagram  of  a  cross  section  of  a  frond  showing  the  distri- 
bution of  an  endophytic  alga  (probably  Chlorochytriuni) .  Larger 
number  on  the  upper  surface,  end.  endophyte. 

Figs.  26  and  27.  Cross  section  of  a  frond ;  detail  drawing  of  endo- 
phyte. iv.  endophyte  wall.  pd.  pyrenoid.  n.  cell  of  nurse  plant  in 
'outline.  Camera  lucida.  x  250. 

BIBLIOGRAPHY. 

1.  Gmelin,  S.  G.     Historia  Fucorum.      1768. 

2.  Postels,  A.,  and  Ruprecht,  F.     Illustrationes  Algarum  in  itinere 
circa  orbem  jussu  Imperatoris  Nicolai.     I.    i8,^/.    ^p,  f.  88.     1840. 

3.  Zanardini,  G.     Saggio  di  classificazione  naturale  delle  Ficee  con 
nuovi  Studi  sopra  1'Androsace  degli  Antichi.  49.    1843. 

4.  Kiitzing,  F.  T.     Phycologia  Generalis.     400.      1843. 

5.  Kiitzing,  F.  T.     Species  Algarum.     744.      1849. 

6.  Agardh,J.    Species,  Genera  etOrdines  Algarum.    2:295.  1851. 

7.  Harvey,  W.  H.     Nereis  Boreali- Americana  i:  21  ;  2:  160,  173. 

'853- 

8.  Harvey,  W.  H.     Notice  of  a  Collection  of  Algae  made  on  the 
Northwest  Coast  of  North  America,  chiefly  at  Vancouver's  Island,  by 
David  Lyall,   Esq.,  M.D.,    R.N.,    in   the   years    1859-61.      Journ. 
Linn.  Soc.  Bot.  6:  172.      1862. 

9.  Kiitzing,  F.  T.     Tabula-  Phycologicas  17:  24.  pi.  83.      1867. 

10.  Zanardini,  G.      Iconographia  Phycologica  Adriatica  2 :  pi.  78. 
1860-76. 

11.  Agardh,    J.     Species,   Genera    et  Ordines  Algarum.    3:   225. 
1876. 


190 


MINNESOTA    BOTANICAL    STUDIES. 


12.  Bornet,  E.     Algues  de  Madagascar  recoltees  par  M.  Ch.  Thie- 
baut.  Bull.  Soc.  Bot.  de  France.  32:  iS./.  i,  2.      1885. 

13.  Hauck,  F.     Die  Meeresalgen,   in  Rabenhorst's  Krypt.  Fl.  2: 
146.  1885. 

14.  Schmitz,  F.     Systematische   Ubersicht    der   bisher   bekannten 
Gattungen  der  Florideen.     Flora  72  :  435-456.      1889. 

15.  Okamura,  R.     Remarks  on  some  algae  from  Hokkaido.     Bot. 

Mag.  Tokyo  5  :  333-336-      l89J  • 

1 6.  Schmitz,  F.,  and  Hauptfleisch,  P.      Die    natiirlichen  Pflanzen- 
familien.     I.  Teil.  Abt.  2.   517,  519,  520,  525.      1897. 


OL.   II. 


MINNESOTA    I 


PL 


NICAL    STUDIES. 


PART 


14 


(U 


t\ 


13 


;  xvn. 


MINNESOTA    BOTANICAL    STUDIES. 


PART    II. 


PLATE    XVIII 


XIV.     EXTENSION     OF     PLANT    RANGES    IN   THE 
UPPER  MINNESOTA  VALLEY. 


L.  R.  MOVER. 


The  following  notes  refer  to  plants  that  have  been  collected 
in  the  upper  Minnesota  valley  since  the  publication  of  Professor 
MacMillan's  Metaspermce  of  the  Minnesota  Valley.  Duplicates 
have  been  deposited  in  the  Herbarium  at  the  State  University. 

Thlaspi  arvense  L. 

This  old  world  crucifer  has  become  well  established  in  the 
railroad  yards  at  Montevideo  and  is  spreading  rapidly. 

Conringia  orientalis  (L,.}  DUMORT. 

This  plant,  first  collected  in  wheat  fields  in  Chippewa  County 
in  1894,  is  spreading  very  rapidly  and  seems  likely  to  become 
one  of  the  worst  "  mustards  "  with  which  the  farmer  has  to  con- 
tend. 

Sisymbrium  altissimum  L. 

This  plant,  too,  is  spreading  very  rapidly  along  the  railroad 
tracks  at  Montevideo,  and  seems  likely  to  become  a  very  trouble- 
some weed. 

Peucedanum  nudicaule  (PURSH)  NUTT. 

This  plant  is  found  to  be  quite  plentiful  in  the  upper  Minne- 
sota valley  on  thin  gravelly  land  near  ledges  of  igneous  rock. 
It  is  one  of  the  earliest  spring  flowers. 

Potentilla  hippiana  LEHM. 

This  western  Potentilla  is  occasionally  found  on  ledges  of 
gneiss  rock  near  Montevideo. 

Helianthus  annuus  L. 

Among  the  recent  arrivals  at  Montevideo,  an  immigrant  from 
the  West,  is  the  annual  sunflower.  It  is  traveling  along  the 
railroads. 


192  MINNESOTA    BOTANICAL    STUDIES. 

Grindelia  squarrosa  (PURSH)  DUNAL. 

This  is  another  western  plant  that  seems  to  be  traveling  east. 
It  has  but  recently  become  well  established  in  the  railroad  yards 
at  Montevideo. 

Lactuca  scariola  L. 

Another  Old  World  weed  that  has  but  recently  arrived  at 
Montevideo  is  this  species.  It  is  spreading  with  great  rapidity. 

Senecio  vulgaris  L. 

Recently  arrived  at  Montevideo,  this  plant  is  becoming  com- 
mon as  a  weed  in  gardens  and  waste  grounds. 

Poa  pseudopratensis  SCRIB.  &  RYD. 

A  Poa  collected  at  Clara  City,  in  Chippewa  County,  has  been 
identified  by  Professor  Lamson-Scribner  as  this  species. 

Fraxinus  lanceolata  BORCK. 

A  study  of  this  species  based  on  a  large  collection  of  ma- 
terial from  the  prairie  portion  of  the  State,  some  of  which  was 
sent  east  for  comparison,  makes  it  probable  that  all  of  our  ash 
trees  are  referable  to  this  species.  F.  Americana  appears  to  be 
absent  from  western  Minnesota. 

Cactus  viviparus  NUTT. 

This  cactus  is  found  quite  abundantly  on  granite  ledges  in  the 
ancient  valley  of  the  "Warren,"  about  two  miles  southeast  of 
Ortonville.  Its  bright  red  flowers  are  strikingly  beautiful. 

Astragalus  flexuosus  (HOOK.)  DOUG. 

This  species  is  quite  plentiful  near  the  railroad  yards  at  Or- 
tonville. 


XV.     LIST   OF   HEPATIC.'E   COLLECTED   ALONG 

THE  INTERNATIONAL   BOUNDARY    BY 

J.  M.   HOLZINGER,   1897. 


ALEXANDER  W.  EVANS. 


1.  Aplozia  autumnalis  (DC.)  HEEG.     F,  N. 

2.  Bazzania  trilobata  (L.)  S.  F.  GRAY.     F,  R,  U. 

3.  Blepharostoma  trichophyllum  (L.)  DUMORT.     R. 

4.  Cephalozia  catenulata  (HUBEN.)  SPRUCE.     F. 

5.  C.  media  LIXDB.     P. 

6.  Frullania  Eboracensis  GOTTSCHE.     F. 

7.  Jungermannia  barbata  SCHREB.     F,  G,  P. 

8.  J.  quinquedentata  WEB.     P,  R. 

9.  J.  ventricosa  DICKS.     S. 

10.  Lejeunea  serpyllifolia  (DICKS.)  LIB.     U. 

11.  Lepidozia  reptans  (L.)  DUMORT.     P,  R,  S,  U. 

12.  Plagiochila  asplenioides  (L.)  DUMORT.     N,  P,  U. 

13.  Porella  platyphylla  (L.)  LINDB.     F,  U. 

14.  Ptilidium  ciliare  (L.)  NEES.     C,  F,  S,  U. 

15.  Radula  complanata  (L.)  DUMORT.     C,  F,  U. 

1 6.  Scapania  glaucocephala  (TAYL.)  AUST.     F. 

C  =  Camp  IV.,  on  the  Prairie  Portage,  shore  of  Basswood 
Lake,  near  the  rapids  from  Sucker  Lake. 

F  =  Fall  Lake,  near  the  foot  of  Kawasatchong  Falls,  seven 
miles  north  of  Ely. 

G  =  between  Gunflint  Lake  and  Grand  Portage. 

N  =  stream  flowing  from  North  Lake  into  Little  Gunflint 
Lake. 

P  =  Grand  Portage  Island. 

R  =  Pipestone  Rapids,  on  Basswood  Lake. 

S  =  Safety  Island. 

U  =  United  States  Peninsula. 


XVI.     OBSERVATIONS    ON   CHLOROCHYTRIUM. 


E.  M.  FREEMAN. 


In  1850  Mettenius  found  numerous  green  cells  in  the  thallus 
of  Polyides  lumbricalis  which  resembled  closely  what  are 
now  classified  as  the  Endosphasreae  of  the  Protococcaceae.  He 
interpreted  them  as  spore-mother  cells  of  the  red  seaweed 
upon  which  they  were  found.  Thuret  fourteen  years  later 
observed  these  same  structures  and  interpreted  them  as  parasitic 
zoospores  which  on  germination  produce  the  bushy  thallus  of 
Cladophora  lanosa.  Cohn  in  1865  was  able  to  confirm  the 
observations  of  Mettenius  and  of  Thuret,  except  as  to  the  germ- 
ination of  the  endophyte  into  Cladophora  lanosa. 

The  condition  of  endophytism  was  considered  at  that  time  as 
indicative  of  parasitism.  Hence  new  interest  was  aroused  in 
the  investigation  of  these  lower  forms  when  Rees  and  Schwen- 
dener  at  about  the  same  time  (1871)  advanced  independently 
the  theory  that  the  Cottema  type  of  lichen  is  to  be  derived  from 
a  discomycetous  fungus,  the  mycelium  of  which  has  pene- 
trated the  mucilage  of  a  Nostoc  completely  surrounding  the 
latter.  Reinke's  observations  on  Nostoc  in  the  stems  of  Gun- 
ner a  scabra  and  the  work  of  Milde  and  Janczewski  on  Nostocs 
in  liverworts  demonstrated  the  occurrence  of  Protococcus-like 
algal  forms  in  the  plant  tissues  of  higher  plants.  Cohn  in 
1872  suggested  that  the  presence  of  the  Nostoc  filaments  in 
Gunner  a  and  Anthoceros  is  perhaps  to  be  explained  as  an  acci- 
dental entry  of  the  movable  Nostoc  filaments  into  the  tissues  of 
the  nurse  plant,  their  continued  growth  in  this  new  sheltered 
position  and  their  subsequent  imprisonment  by  the  growth  of 
the  surrounding  tissues  of  the  nurse  plant.  In  contrast  to  this 
form  of  endophytism  Cohn  describes  the  new  genus  Chloro- 
chytrium,  which  he  considers  to  be  a  true  parasite  in  certain 
species  of  Lemna.  The  zoospores,  very  numerous  on  the  sur- 
face of  the  host,  send  out  a  germination  tube  between  two  epi- 

195 


196  MINNESOTA    BOTANICAL    STUDIES. 

dermal  cells.  The  membrane  of  this  tube  becomes  thickened 
by  subsequent  layers,  the  tube  swells  with  the  absorption  of  the 
chlorophyll  and  protoplasm  and  the  intercellular  endophyte 
results,  with  a  cellulose  button  protruding  from  the  point  where 
the  germinating  tube  entered.  The  endophyte  then  becomes 
pyriform  and  almost  opaque  on  account  of  the  density  of  the 
chlorophyll.  By  free  cell  formation  large  cells  are  formed  in 
the  endophyte  and  these  finally  break  into  a  large  number  of 
zoospores  which  are  expelled  through  the  cellulose  protuber- 
ance from  the  nurse-plant  epidermis.  The  endophyte  is,  there- 
fore, an  independent  organism  closely  related  to  Hydrocytium 
(Characium  A.  Br. )  on  the  one  hand  and  to  Synchytrium  on 
the  other.  With  the  Eu-Synchytrium  group  its  cell  form  and 
the  formation  of  zoospores  by  a  preliminary  division  into  seg- 
ments, corresponding  to  the  zoosporangia  of  Synchytrium, 
agree,  but  it  differs  in  the  presence  of  chlorophyll  and  of  a 
germination  tube  and  in  its  intercellular  position.  Upon  these 
observations  Cohn  founded  the  genus  Chlorochytrium  and  de- 
scribed it  as  follows  : 

Planta  endophytica  viridis  unicellaris,  globosa  ovoidea  vel 
irregulariter  curvata  bi,  tri,  multiloba  dense  conferta  plasmate 
viridi,  primum  in  segmenta  majora  diviso  dein  secedente  in 
zoosporas  immersas  pyriformes  virides  processibus  tubulosis  ex- 
tus  emissas. 

Chlorochytrium  lemna  upon  which  the  genus  is  based  is  then 
described. 

Cohn  pronounces  Chlorochytrium  a  true  parasite.  That  no 
deleterious  effects  upon  the  host  are  visible  is  paralleled  in  Per- 
onospora  and  Synchytrium.  In  its  intercellular  position  it  re- 
sembles the  Uredineae. 

Two  years  after  Cohn's  observations  were  published  Kny 
described  a  new  species  of  Chlorochytrium  endophytic  upon 
Ceratophyllum  demersum.  It  differs  from  Chlorochytrium 
lemnce  in  size  and  in  the  absence  of  a  cellulose  button. 

In  1877  Wright  established  a  third  species  of  Chlorochy- 
trium, C.  cohnii  Wright. 

"  The  zoospores  impinging  on  the  fronds  of  several  species 
of  marine  algae  quickly  assuming  a  figure-of-eight  form,  the 
lower  sphere  growing  into  the  frond  and  rapidly  assuming  com- 
paratively large  dimensions,  the  upper  sphere  remaining  as  a 
tube-like  neck  portion  to  the  larger  mass.  On  the  cell  arriving 


Freeman  :    OBSERVATIONS  ON  CIILOROCHYTRIUM.  197 

at  an  adult  stage,  the  whole  of  the  green  protoplasmic  contents 
divides  into  a  number  of  from  10—30  nearly  circular  zoospores, 
which  escape  through  the  neck-shaped  portion. 

"  Living  in  the  thallus  of  various  species  of  Schizonema, 
Polysiphonia,  etc.  ;  also  on  the  Infusoria  found  at  Howth." 

Wright  states  further  that  there  are  two  kinds  of  zoospores, 
large  and  small,  the  latter  being  the  more  numerous. 

Szymanski  in  1878  described  C.  knyanum  apparently  identical 
with  the  plant  mentioned  by  Kny  four  years  before  as  inhabiting 
the  tissues  of  Ceratophyllum  demersum.  This  species  was  found 
on  Lcmna  minor  and  possessed  a  cellulose  button  which  did  not 
protrude  farther  than  twice  the  thickness  of  its  outer  wall  above 
the  epidermis  of  its  nurse  plant. 

Klebs  published  the  results  of  his  investigations  on  C.  lemnce 
in  1881.  Chlorochytrium  in  the  younger  vegetative  stages 
contains  a  light  green  chlorophyll-bearing  protoplasm  with  iso- 
lated starch  grains  surrounded  by  a  cell  sap  vacuole  (see  below, 
pyrenoids) .  In  the  later  stages  the  grains  increase  in  number,  the 
mesh-work  of  green  bands  becomes  smaller,  the  chlorophyll 
darker  until  almost  opaque  and  the  protoplasm  finally  becomes 
coarsely  granular.  After  a  resting  period  of  a  week  or  more 
the  zoospores  are  formed  by  successive  bipartitions  of  the  cell 
contents,  at  first  by  perpendicular,  later  by  radially  disposed 
walls. 

The  number  of  divisions  is  not  known.  Liberation  of  the 
zoospores  is  accomplished  by  absorption  of  water  resulting  in 
the  splitting  of  the  Chlorochytrium  wall  and  of  the  superposed 
Lcmna  tissues. 

The  conjugation  of  the  biciliate  zoogametes  into  larger 
quadriciliate  zoozygotes  was  observed,  a  fact  which  niay  throw 
light  on  the  macro-  and  microzoospores  of  Wright's  species. 
Klebs  observes  that  Cohn's  account  of  the  liberation  of  the 
zoospores  is  without  observational  foundation  and  doubts  its  ac- 
curacy. He  also  calls  in  question  the  appropriateness  of  placing 
Wright's  species  in  the  genus  Chlorochytrium  and  further  sug- 
gests the  probability  that  C.  knyanum  is  but  the  asexual  form 
of  C.  lemncg  since  no  copulation  had  been  observed  between 
the  zoospores.  C.  -pallidum  Klebs  and  many  similar  forms  are 
probably  mere  "  place  varieties"  of  C.  knyanum. 

Klebs  points  out  with  much  truth  that  no  proof  has  been  given 
of  the  much  averred  parasitism  of  Chlorochytrium  by  Colin 


193  MINNESOTA    BOTANICAL    STUDIES. 

and  other  previous  investigators.  On  the  other  hand  Chloro- 
chytrium  has  well  developed  chlorophyll  and  lives  near  the  sur- 
face where  abundant  light  is  available.  The  requisite  inorganic 
matter  may  gain  access  to  the  cells  by  the  constant  or  at  least 
periodical  submersion  in  water. 

Chlorochytrium  lemntz  penetrates  dead  as  well  as  living 
leaves  and  culture  methods  demonstrate  an  entire  lack  of  de- 
pendence of  the  endophyte  upon  a  host  plant.  In  many  endo- 
phytes  zoospores  can  be  developed  on  culture  slides  for  months. 
No  proof  has  as  yet  been  adduced  for  any  injury  of  the  host 
beyond  the  results  of  the  mechanical  pressure  exerted.  The 
explanation  then  of  endophytism  is  to  be  found  not  in  parasitism 
but  in  the  mechanical  protection  of  position,  which  the  inter- 
cellular spaces  of  the  host  offered ;  hence  the  appropriateness 
of  the  term  "  Raum  Parasiten."  It  is,  of  course,  possible  for 
parasitism  to  develop  from  such  a  condition  and  this  develop- 
ment seems  to  be  in  evident  progress  in  such  a  nearly  related 
form  as  Phyttobium  dimorphum  and  also  perhaps  in  Nostoc 
lichenoides.  In  the  systematic  relations  of  Chlorochytrium  and 
the  nearly  related  genera,  Klebs  briefly  points  out  the  inter- 
mediate position  of  Chlorochytrium  and  Endos-ph<Rra>  between 
the  Protococcaceae  and  the  Chytrideee,  the  isolated  position  of 
Scotinosphara  and  the  probable  affinities  of  Phyllobiutn  on 
the  one  hand  with  Chlorochytrium  and  on  the  other  with  Bo- 
try  dium. 

Schaarschmidt,  1881,  found  zoospores  of  Chlorochytrium 
in  a  Desmid  culture  in  which  the  zoospores  subsequently  devel- 
oped, confirming  Kleb's  view  on  the  parasitism  of  the  endo- 
phyte. In  1883  Kjellman  described  the  following  species  : 

Chlorochytrium  inclusum  Kjellman. — "In  the  vegetative 
stage  spherical  or  subspherical,  entirely  included  within  the 
nurse  plant,  with  the  formation  of  the  zoospores  becoming 
slightly  elongated,  short-conical,  flask-shaped,  ovoid  or  ellip- 
soidal, finally  bare  at  the  pointed  apex,  which  penetrates  the  cor- 
tical layer  of  the  nurse  plant  and  emitting  the  zoospores  through 
an  ostiole  which  has  been  formed."  This  species  is  endophytic 
upon  Sarc&phyllis  arctica^  mostly  near  the  surface  but  some- 
times in  the  middle  of  the  host.  It  averages  80-100  mic.,  has 
yellowish-green  contents  and  a  cell-wall  which  is  thin  and  of 
equal  thickness.  The  chromatophore  is  thin  and  is  spread  along 
the  wall.  With  the  elongation  at  the  formation  of  zoospores 


Freeman:    OBSERVATIONS  ON  CHLOROCHYTRIUM.  199 

the  membrane  thickens  towards  the  outer  surface  especially, 
and  a  cone-shaped  growth  of  cellulose  is  formed.  The  proto- 
plasm then  takes  on  a  more  intense  yellow  green  and  divides 
into  numerous  zoospores.  Numerous  bulgings  of  the  plant  cell 
are  produced  probably  by  the  growth  of  the  surrounding  tissues 
of  the  host.  An  ostiole  is  formed  at  the  apex  of  the  cellulose 
out-growth  by  which  the  zoospores  escape.  Those  cells  found 
in  the  center  of  the  host  may  attain  as  great  a  diameter  as  275 
mic.  The  even  thickness  of  the  wall  of  these  cells  suggests 
that  they  may  be  resting  stages.  Kjellman  refers  this  plant  to 
the  genus  Chlorochytrium,  but  hesitatingly  on  account  of  his 
lack  of  knowledge  about  the  further  development.  He  found 
C.  inclusion  in  all  of  the  Sarcophyllis  edulis  material  which  he 
has  examined.  Zoospores  are  most  abundantly  produced  in 
winter,  but  are  also  found  in  summer.  The  endophyte  occurs 
in  greatest  abundance  and  most  strongly  developed  in  Sarco- 
phyllis arctica.  Its  range  is  apparently  coextensive  with  that  of 
Sarcophyllis  arctica;  i.  <?.,  throughout  the  arctic  region,  except 
in  the  North  Atlantic,  most  abundant  in  the  eastern  part  of  the 
Siberian  sea. 

Three  more  new  species  were  described  by  Schroeter  in  1883. 
Chlorochytrium  rubrum  with  red  contents  and  occupying  the  air 
spaces  of  the  leaves  and  stems  of  Pcplis  portula  and  of  Mentha 
aquatica ;  Chlorochytrium  viride,  in  the  leaves  of  Rumex  obtus- 
ifolius;  and  C.  latum,  a  spherical  cell  with  yellow  contents 
which  become  green  in  water,  found  in  Lychnis  jlos-cuculi. 

The  investigations  of  Moore  on  Chlorochytrium  lemna  pub- 
lished in  the  following  year  brought  to  light  no  new  facts  of 
importance.  Moore  held  that  the  nearest  affinities  of  Chlorochy- 
trium lie  with  Protomyces. 

In  1887  Hieronymus  described  C.  archerianum  in  punctured 
cells  of  Sphagnum  leaves,  and  characterized  by  a  greatly  devel- 
oped cellulose  button.  Zoospores  are  formed  regularly  but  do 
not  copulate.  P.  Hariot,  1889,  collected  C.  inclusum  on  species 
of  Gigartina  at  Cape  Horn,  supposedly  identical  with  Kjell- 
man's  C,  inclusum. 

In  his  Conspectus  of  Endophytic  Algse,  Mobius,  1891,  men- 
tions in  addition  to  those  species  enumerated  above,  C.  dermato- 
colax  which  was  described  by  Reinke  and  found  on  species  of 
Polysiphonia  and  Sphacelaria,  and  in  his  opinion  should  be 
classified  under  the  genus  Chlorocystis,  since  it  is  marine,  is  in- 
tracellular  and  emits  zoospores  singly. 


200  MINNESOTA    BOTANICAL    STUDIES. 

C.  schmitzii  was  described  (1893)  by  Rosenvinge  from 
Greenland  material  on  Cruoria  arctica.  The  cell  is  without 
a  cellulose  button  or  papilla,  is  more  or  less  attenuate  at  the 
base  and  has  a  single  chromatophore  with  sometimes  two  pyre- 
noids.  Zoospores  were  not  observed. 

Collections,  Methods,  etc. — Upon  the  Constantinea  material 
which  was  collected  by  Miss  J.  E.  Tilden  at  different  points 
along  Puget  sound  and  was  distributed  as  Constantinea  sitch- 
ensis  Post,  and  Rupr.,  were  found  a  large  number  of  endo- 
phytic  unicellular,  chlorophyllaceous  algas,  referred  to  the 
genus  Chlorochytrium  of  Cohn.  The  endophyte  was  found  in 
abundance  on  all  of  the  Constantinea  collected.  One  collection 
was  made  in  August,  1897,  near  Seattle,  and  in  the  summer  of 
1898  (May  25-July  2),  five  were  made  at  as  many  points  farther 
up  the  Sound.  Most  of  the  material  used  in  the  following  in- 
vestigations was  fixed  and  preserved  in  a  2  per  cent,  sea-water 
solution  of  formalin.  The  green  color  of  the  endophyte  was 
well  preserved.  The  lower  ends  of  the  cells  have  in  very  many 
cases  an  irregular  outline  which  may  possibly  be  in  part  due  to 
shrinkage  but  is  caused  for  the  most  part  by  pressure  of  the 
surrounding  Constantinea  cells.  In  all  of  this  endophyte 
formalin  material  and  in  the  dried  material  as  well,  though  not 
so  markedly,  brown  bodies  were  found  jutting  out  between  the 
chromatophore  and  the  cell  wall  and  assuming  various  forms 
(see  below).  These  bodies  were  undoubtedly  due  to  a  chloro- 
phyllan  reaction,  the  hypochlorin  reaction  of  Pringsheim.  The 
formalin  solution  when  tested  was  found  to  give  a  slightly  acid 
reaction. 

Sections  of  Constantinea  cut  freehand  between  elder  pith 
furnished  most  of  the  material  for  study.  They  were  mounted 
either  in  the  two  per  cent,  formalin  sea-water  solution,  in  glyc- 
erine, or  in  glycerine  jelly.  Material  carried  through  the  usual 
paraffin  method  stained  and  mounted  in  Canada  balsam  has  also 
furnished  useful  sections.  The  abundance  of  the  endophyte 
makes  it  easy  to  get  favorable  surface  and  sectional  views  of  it. 

General  Habit  and  Structure. — The  endophytes  on  Constan- 
tinea sitchensis  occur  on  both  the  upper  and  lower  surfaces  of 
the  fronds.  I  have  been  unable  to  find  any  on  the  stipe.  They 
are  most  abundant  on  the  older  fronds  and  especially  towards 
the  peripheral  portion.  From  some  young  fronds  they  are 
almost  altogether  wanting.  They  occur  in  greatest  numbers 


Freeman  :    OBSERVATIONS  ON  CHLOROCHYTRIUM.  201 

on  the  lower  surface  while  rather  few  are  found  on  the  upper 
side.  The  following  figures  are  taken  from  a  medium-sized 
frond  in  the  peripheral  region :  on  the  under  side  140-160 
(sometimes  as  many  as  230)  in  one  square  mm.  ;  on  the  upper 
surface  60-65  m  tne  same  area.  Many  areas  of  4  square  mm. 
on  the  upper  surface  contained  no  endophytes. 

The  endophytic  cells  are  found  almost  exclusively  in  the  tis- 
sues just  beneath  the  pseudoepidermis  of  the  nurse  plant,  with 
the  slightly  pointed  end  just  at  or  just  below  the  surface.  They 
occur  in  rare  cases  in  the  central  part  of  the  frond  completely 
enclosed.  The  pointed  e.id  protrudes  from  the  tissues  of  the  nurse 
plant  in  but  few  cases  and  then  not  mor-j  than  for  a  distance 
equal  to  one-half  the  thickness  of  its  outer  wall.  The  cells  not 
infrequently  occur  between  the  paraphyses  of  the  nemathecia 
of  Constantinea  where  they  usually  penetrate  to  but  not  into  the 
tissues  beneath. 

The  predominant  form  assumed  by  the  endophyte  is  pear- 
shaped  with  the  smaller  end  directed  toward  the  surface  of  the 
nurse  plant.  The  cells  are  often  ovoidal  and  even  ellipsoidal. 
In  the  central  portion  they  assume  a  spheroidal  form.  In  the 
paraphyses  they  become  elongated  or  assume  a  figure-of-eight 
form  similar  to  that  described  by  Wright  for  Chlorochytrium 
cohnii.  The  inner  ends  of  the  cells  are  marked  more  or  less 
by  the  bulgings  undoubtedly  caused  by  the  pressure  of  the 
adjacent  cells  of  the  nurse  plant. 

The  cells  average  85-115  mic.  in  length  and  40-85  mic.  in 
breadth  but  often  attain  143  x  100  mic.  The  wall  in  some  cases 
is  28  mic.  thick  at  the  outer  surface  and  8  mic.  thick  around  the 
remainder  of  the  cell,  but  usually  is  less  than  one-half  of  these 
dimensions.  The  lamellation  of  the  cell  wall  can  clearly  be 
seen  in  many  sections  (especially  glycerine  mounts)  and  is  due 
probably  to  the  apposition  of  successive  layers  of  cellulose. 
Chlorophyll  occurs  in  the  form  of  a  single  yellowish-green  plate 
in  which  are  included  a  large  number  of  fine  refringent  gran- 
ules. This  chromatophore  extends  around  the  entire  wall  of 
the  cell  and  contains  a  varying  number  of  very  conspicuous 
pyrenoids,  which  are  flattened  spherical  in  form,  5-11  mic.  in 
diameter,  and  jut  out  into  the  cavity  of  the  cell.  As  many  as 
thirty-nine  have  been  found  in  a  single  cell  and  at  least  one 
pyrenoid  can  be  seen  soon  after  the  cell  begins  to  penetrate  the 
tissues  of  the  nurse  plant.  The  pyrenoids  show  a  clear  central 


202  MINNESOTA    BOTANICAL    STUDIES. 

portion  probably  proteid.  In  sections  stained  heavily  with 
aniline  blue  the  central  portion  appears  blue.  Around  the  clear 
center  are  arranged  5-10  plates  of  starch  which  stain  brown 
with  both  a  KI  and  an  alcohol  solution  of  iodine.  By  careful 
washing  of  material  stained  in  an  alcoholic  solution  of  iodine 
and  with  the  aid  of  a  -^  oil  immersion  lens  a  distinct  violet 
tinge  is  discernible  in  the  plates.  The  protoplasmic  contents  of 
the  cell  are  usually  most  dense  in  the  pointed  part.  Between 
the  chromatophore  and  the  cell  wall  are  found  numerous 
rust-brown  to  black  (in  a  few  cases  copper-colored)  bodies 
of  different  form  and  size.  In  some  places  they  occur  in  diffuse 
patches  the  limits  of  which  are  often  indefinable,  in  others  as 
five-pointed  rosettes.  Again  they  may  appear  filiform,  partially 
and  usually  irregularly  coiled  or  forming  a  delicate  and  loose 
network.  I  have  interpreted  these  bodies  as  products  of  the 
action  of  the  dilutely  acid  formalin  solution  and  as  identical 
with  the  hypochlorin  of  Pringsheim.  His  plates  agree  closely 
with  much  of  the  material  at  hand.  In  accordance  with 
Pringsheim's  account  of  the  chemical  reactions  of  hypochlorin, 
these  brown  bodies  are  wanting  in  those  sections  which  have 
been  carried  through  the  alcohols  in  the  method  for  paraffin 
embedding. 

A  large  amount  of  material  has  been  examined  but  in  no 
case  has  even  a  trace  of  the  production  of  zoospores  or  gametes 
been  found.  The  stages  in  the  penetration  of  the  nurse  plant, 
consisting  in  the  elongation  of  the  at  first  spheroidal  cell,  the 
subsequent  withdrawal  of  the  protoplasm  into  the  inner  end  and 
the  increase  in  size  of  the  latter  to  form  the  mature  pyriform 
cell,  have  been  observed,  but  nothing  to  indicate  the  formation 
of  zoospores. 

Conclusions. — It  is  therefore  upon  the  basis  of  vegetative 
characters  that  the  endophyte  described  above  is  provisionally 
placed  in  the  genus  Chlorochytrium  under  C.  inclusum  Kjell- 
man.  Upon  examination  of  the  Chlorochytrium  inclusum  found 
upon  Dilsea  (Sarcophyllis}  distributed  in  Phycotheca  Boreali- 
Americana  (Fasc.  XL,  no.  514)  this  is  seen  to  possess  a 
thicker  cell  wall  than  the  material  on  Constantinea  sitchensis,  is 
almost  spheroidal,  larger,  has  denser  dark  green  contents,  con- 
tains no  pyrenoids  (or  very  inconspicuous,  if  present  at  all)  and 
resembles  a  resting  stage.  The  time  of  collection,  December, 
moreover,  strengthens  this  last  supposition.  The  material  under 


Freeman  :    OBSERVATIONS  ON  CHLOROCHYTRIUM.  203 

observation  was  collected,  on  the  other  hand,  in  summer,  May 
to  July.  The  light  yellow-green  color,  the  absence  of  repro- 
ductive bodies  and  the  abundance  of  small  cells  point  strongly, 
I  think,  to  the  conclusion  that  this  endophyte  is  but  a  young 
stage  of  Chlorochytrium  inclusum  Kjellman.  A  similar  dif- 
ference in  the  vegetative  and  resting  stages  of  C.  lemnce  are 
recorded  in  Klebs'  observations  (pi.  39,  f.  2  and  p).  The 
form  and  habit  of  the  endophyte  upon  Constantinea  accord 
best  with  Kjellman's  description  of  C.  inclusum\  no  mention, 
however,  is  made  by  Kjellman  of  pyrenoids,  which  are  so 
conspicuous  in  the  Constantinea  material.  It  is  suggested  by 
De-Toni  and  also  by  Miss  Whitting  that  Kjellman's  species 
may  possibly  belong  to  Chlorocystis,  a  genus  established  by 
Reinhard  in  1885.  Chlorocystis  is  described  with  but  one 
pyrenoid  while  the  endophyte  on  Constantinea  contains  many. 
C.  schmitzii,  described  by  Rosenvinge,  is  but  imperfectly 
known.  I  have  seen  in  the  sections  studied  a  number  of  cases 
where  the  endophyte  in  Constantinea  has  assumed  approxi- 
mately the  same  irregular  or  obovoidal  form  shown  in  the 
figures  of  Rosenvinge.  The  only  remaining  marine  species 
of  Chlorochytrium,  C.  dermatocolax,  lives  in  the  outer  mem- 
brane of  Polysiphonia  and,  according  to  Mobius,  belongs  prob- 
ably to  Chlorocystis. 

The  great  similarity  in  vegetative  structure  to  that  described 
by  Kjellman  for  C.  inclusum  would  indicate  that  the  Constan- 
tinea endophyte  described  above  is  a  midsummer  stage  of  C. 
inchisum  and  I  would  provisionally  place  it  in  that  species 
awaiting  further  information  on  the  life  history  and  develop- 
ment. 

BIBLIOGRAPHY. 

Cohn,  F.  Ueber  parasitische  Algen.  Remarks  before  the  "  Wand- 
erversammlung  der  Schlesischen  Gesellschaft  fur  vaterlandischer  Cul- 
tur.  Botanische  Section.  1872.  Beit,  zur  Biologic  der  Pflanzen. 
I1:  87.  pi.  2.  1875. 

Kny,  L.  Ueber  eine  griine  parasitische  Alge.  Sitz.  Gesellch.  na- 
ttirf.  Freunde  zu  Berlin.  1874. 

Wright,  P.  On  a  new  species  of  parasitic  green  alga  belonging  to 
the  genus  Chlorochytrium  of  Cohn.  Trans.  Roy.  Irish  Acad.  26:  1877. 

Szymanski,  F.  Ueber  einige  parasitische  Algen.  Inaug.  dissert- 
der  Univ.  Breslau.  1878. 

Kirchner,  O.     Die  Algen  Schlesiens.     Breslau.      1878. 

Klebs,  G.  Beitrage  zur  Kenntniss  niederer  Algenformen  Bot. 
Centralb.  39:  16-21.  pi.  j,  4.  1881. 


204  MINNESOTA    BOTANICAL    STUDIES. 

Hariot,  P.  Algues  recuillies  par  la  Mission  scientifique  du  Cap 
Horn.  Paris.  1882-83. 

Reinke,  J.  Algenflora  der  westliche  Ostsee  deutschen  Antheils.  Sep.- 
Abdr.  VI.  Ber.  Commiss.  Untersuch.  deutschen  Meere,  in  Kiel.  1889. 

De-Toni,  J.  B.     Sylloge  Algarum.   x:  635-637.      1889. 

Mobius,  M.  Conspectus  Algarum  endophytarum.  Notarisia.  6: 
1282.  1891. 

Rosenvinge,  M.  L.  Kolderup.  Les  Algues  Marines  du  Greenland. 
Ann.  Sci.  Nat.  Bot.  19:  169.  f.  56.  1894. 

Whitting,  F.  G.  On  Chlorocystis  sarcophyci.  Phyc.  Mem.  Pt.  2. 
41-45.  1893. 

Wille,  N.  Die  natiirlichen  Pflanzenfamilien.  I.  Abt.  2.  66.  f. 
42,  1897. 

Schaarschmidt,  G.  A  Chlorochytrium  Erdelylen.  Magyar  N6- 
venyt.  Lapok.  5:  37.  8181. 

Kjellman,  F.  R.  Algae  of  the  Arctic  sea .  320.  pL  31- f.  8-17.   1883. 

Schroter,  J.  Neue  Beitrage  zur  Algenkunde  Schlesiens.  61. 
Jahresb.  Schles.  Gesellch.  Cult.  178-189.  1883. 

Moore,  S.  Le  M.  Remarks  on  some  endophytic  Algae.  Journ. 
of  Bot.  22:  136-138.  1884. 

Lagerheim,  G.  .  On  Chlorochytrium  cohnii  Wright  och  ders  for- 
hallande  till  narstande  arter.  Sv.  V.  A.  Oefvers.  7  :  91-97.  pL  i. 
(Bot.  Jahres.  I21.'37i.  1886.) 

Hieronymus,  G.  Ueber  einige  Algen  des  Riesengebirges.  Jahresb. 
Schles.  Gesellch.  Vaterl.  Cultur.  293-297.  1887. 

Tilden,  J.  E.     American  Algae.     Cent.  III.      1898. 

DESCRIPTION  OF  PLATE  XIX. 

1.  Diagram  of  a  cross  section  of  a   frond  of  Constantinea  sitchen- 
sis  showing  the  endophytic    Chlorochytrium  cells  on  the  upper   (less 
numerous)   and  lower  surface,     e.  endophyte.     p.  pseudoepidermis. 
*'.  intermediate  area.     c.  central  area  of  the  frond. 

2.  Surface  view   of  the    frond    of    Constantinea    (lower  surface) 
showing  the  pseudoepidermis  (/)  with  cavities  (e^  through  which  the 
endophytes  have  penetrated,     x  57. 

3.  4,  5,  6.     Stages  in  the  penetration  of  the  nurse  plant,     s.  pyre- 
noid.     x  288. 

7.  Chlorochytrium  cell  with  a   not  infrequent  form.     Shows  fila- 
mentous form  of  hypochlorin.      x  288. 

8.  Cell  showing  elongated  form  of  endophyte  found  amongst  the 
paraphyses  of  the  nurse  plant,     x  288. 

9.  Large  Chlorochytrium  cell  of  typical  form,     x  288. 

10.  Detail    showing  the   position  of  a  Chlorochytrium  cell  in  the 
cross  section  of  the  Constantinea  frond.      Letters  as  in  no.  i.      x  288. 

n.  The  outer  end  of  a  Chlorochytrium  cell  showing  the  lamella- 
tion  of  the  cell  wall,  x  288. 


VOL.  II.  MINNESOTA  BOTANICAL  STUDIES. 


PART  III. 


10 


PLATE  XIX. 


THE    HELIOTYPE    PRINTING   CO..    BOSTON. 


XVII.     OBSERVATIONS  ON  RHODYMENIA. 


FREDERIC  K.   BUTTERS. 


History  and  Literature. — The  genus  Rhodymenia  was 
founded  by  Greville  in  1830.  As  originally  constituted  it  con- 
tained species  which  have  since  been  referred  to  Gracilaria, 
Gigartina,  Kattymcnia,  Calliblepharis  and  other  genera. 
Agardh  (3),  p.  376,  states  that  in  his  Mediterranean  Algae  (i) 
he  transferred  many  species  to  the  genera  Gracilarja,  Gigan- 
tina  and  Kallymenia,  but  united  Calliblepharis  \\ith  Rhody- 
menia under  the  latter  name.  In  1849  Kiitzing  in  his  Species 
Algarum,  p.  778,  united  the  species  of  the  genera  Rhodymenia 
and  Gracilaria  together  with  some  other  species  under  the 
generic  name  Sphcero  coccus. 

Agardh  (2),  p.  15,  (3)  p.  375,  revised  the  genus  Rhodymenia 
and  gave  it  substantially  its  present  limits  and  generic  characters. 
The  generic  description  as  given  by  De-Toni  and  Levi,  p.  19, 
is  as  follows  :  Frond  plane,  membranaceous,  dichotomous  or  pal- 
mate, proliferate  from  the  margin  or  surface,  composed  of  two 
layers  ;  interior  cells  oblong,  cortical  minute,  rounded  ;  cystocarps 
scattered  throughout  the  frond,  each  within  an  hemispherical 
pericarp  opening  by  a  carpostome,  composed  of  cells,  the  outer 
radiating  the  inner  concentric ;  cystocarp  with  a  simple  rounded 
or  somewhat  lobed  nucleus  ;  nucleus  naked  within  the  pericarp, 
on  a  basal  placenta  with  paniculate-branched  placental  fila- 
ments sustaining  the  lobes  ;  young  fertile  lobes  disposed  radially 
composed  of  articulated  filaments,  older  obconico-rotund,  bear- 
ing numerous  protospores ;  tetraspores  often  collected  into  sori, 
cruciately  divided ;  antheridia  produced  in  superficial  sori  com- 
posed of  minute  hyaline  cells  in  a  single  vertical  series. 

Greville  spelled  the  name  of  this  genus  Rhodomenia.  Mon- 
tagne,  p.  44,  in  1839,  employed  this  spelling  in  a  list  of  Bra- 
silian  cryptogams,  but  in  a  footnote  states  that,  in  conform- 
ity to  its  etymology  it  should  be  spelled  Rhodymenia.  J. 

205 


206  MINNESOTA    BOTANICAL    STUDIES. 

Agardh  (3)  adopted  the  latter  spelling  and  it  has  since  been  in 
general  use,  although  Rhodomcnia  and,  according  to  Ardis- 
sone,  Rhodhymenia  and  Rodkymenia  also  have  been  occasion- 
ally employed. 

Rhodymenia  pertusa  was  first  described  and  figured  by  Pos- 
tels  and  Ruprecht,  p.  20,  pi.  j6,  under  the  name  Porphyra 
pertusa.  Kiitzing,  p.  693,  describes  it  under  this  name.  He 
introduces  a  question,  however,  in  respect  to  its  generic  de- 
termination. J.  G.  Agardh  (3),  p.  376,  points  out  that  the  spe- 
cies should  be  classed  as  a  Rhodymenia  and  not  as  Porphyra. 
He  describes  it  as  Rhodymenia  pertusa  (Post,  and  Rupr.)  and 
places  it,  together  with  Rhodymenia  palmata  (Linn.  Sp.  2  : 
1630)  and  Rhodymenia  peruviana  (J.  Ag.  Mscr.)  in  the  sec- 
tion Palmata  characterized  by  "  tetraspores  occurring  through- 
out the  surface  of  the  frond,  scattered  or  collected  into  sori." 
In  his  Epicrisis  (Agardh,  J.  G.  (4),  p.  379),  he  assigns  it  the 
same  position. 

Agardh  (3)  gives  the  habitat  of  Rhodymenia  pertusa  as  "  in 
the  Arctic  Ocean  near  Kamtschatka  (Mus.  Petropolitani !)  ; 
and  near  Greenland  (Wormskjold  !)."  Kjellman  reports  it 
it  from  the  northwest  coast  of  Spitzbergen  and  the  west  coast  of 
Greenland.  It  was  first  reported  from  the  northwest  coast  of 
America  by  Harvey  (i)  who  found  it  among  the  algae  collected 
by  Captain  Wilkes'  exploring  expedition.  Harvey  gives  its 
locality  as  the  Straits  of  St.  Juan  de  Fuca.  He  compared 
Wilkes'  specimens  with  an  authentic  specimen  furnished  him 
by  Dr.  Ruprecht  and  found  them  identical  in  species,  though 
Wilkes'  specimens  were  considerably  larger  than  Ruprechts's. 
Harvey,  p.  171,  also  reports  the  species  as  collected  by  Dr. 
Lyall  in  1859-61,  his  specimens  being  "  cast  ashore  on  Point 
Roberts,  and  on  rocks  at  low  water,  Fuca  Strait."  Cystocarps 
were  present  on  both  sets  of  specimens  mentioned  by  Harvey. 
In  commenting  on  the  specific  name,  Harvey  (i)  p.  148,  states 
that  to  him  the  perforations  of  the  thallus  appear  to  be  due  to 
casualties. 

In  1893  Carruthers,  p.  80,  examined  one  of  the  specimens 
mentioned  by  Harvey  as  collected  by  Dr.  Lyall.  He  found 
that  in  that  specimen  the  cystocarps  occur  all  over  the  much 
perforated  segments ;  the  majority  are  very  young  and  project 
but  little  from  the  surface  of  the  thallus.  The  structure  of  the 
thallus  is  that  typical  of  the  genus.  He  says  "  The  cystocarp 


Butters:   OBSERVATIONS  ON  RHODYMENIA.  207 

projects  on  one  side  of  the  thallus  and  possesses  a  fairly  thick 
fruit  wall  of  five  or  six  cells  in  thickness  arranged  irregularly 
inwards.  At  the  bottom  of  the  fruit  cavity  is  to  be  found  the 
placenta  formed  of  numerous  small  cells  closely  packed  to- 
gether and  lying  on  the  medullary  layer  of  large  cells. 

"  From  the  placenta  the  gonimoblast  of  more  or  less  irregular 
shape,  is  borne  on  an  elongated  style  cell  and  spreads  upward 
into  the  empty  fruit  cavity. 

"This  gonimoblast  is  composed  of  numerous  pear-shaped 
lobes  which  lie  close  together.  The  spores  of  these  lobes  be- 
come ripe  nearly  at  the  same  time. 

"  The  ostiole  is  generally  quite  in  the  center  of  the  projecting 
fruit  wall,  and  is  similar  to  the  ostioles  of  the  kindred  species." 

Collection  and  Preservation. — All  the  material  at  hand  was 
collected  by  Miss  Josephine  E.  Tilden  at  Port  Orchard,  Kitsap 
County,  Washington,  on  August  2,  1897.  The  specimens  were 
dredged  in  water  4-6  fathoms  deep.  A  small  portion  of  the 
material  was  killed  and  preserved  in  80  per  cent,  alcohol.  The 
larger  part  of  the  material  was  dried.  By  soaking,  this  dried 
material  so  far  regained  its  original  form  that  the  anatomy  could 
be  well  studied  although  the  cell  contents  were  largely  disor- 
ganized. All  the  observations  upon  the  stipe  and  the  prolifera- 
tions were  made  upon  this  dried  material ;  observations  upon 
other  points  were- made  mainly  upon  the  alcoholic  material. 

Methods. — The  dried  material  was  soaked  in  water  until  it 
regained  its  natural  consistency.  Various  methods  were  em- 
ployed in  cutting  the  tissues.  Much  of  the  material  was  cut 
upon  the  Osterhout  freezing  chamber.  (Osterhout,  W.  J.  V., 
p.  1950  The  alcoholic  material  was  first  passed  into  water 
—preferably  through  about  three  intermediate  grades  of  dilute 
alcohol.  When  the  alcohol  was  completely  removed  the  mate- 
rial was  in  some  cases  infiltrated  with  gelatine  solution  and  then 
mounted  in  a  drop  of  gum  arabic  solution  upon  the  freezing 
chamber ;  in  other  cases  it  was  embedded  directly  in  the  gum 
arabic.  On  account  of  the  firm  nature  of  most  of  the  tissues, 
this  method  of  mounting  directly  in  gum  arabic  proved  quite  as 
successful  as  that  in  which  the  tissues  were  first  infiltrated  with 
gelatine. 

The  sections,  as  soon  as  they  were  removed  from  the  knife, 
were  passed  into  20  per  cent,  glycerine.  Those  which  were  to 
be  stained  were  transferred  from  this  glycerine  to  the  staining 


208  MINNESOTA    BOTANICAL    STUDIES. 

solution,  thence,  after  washing,  back  to  the  glycerine  solution. 
The  glycerine  solution  was  allowed  to  concentrate  by  evapora- 
tion, and  the  sections  were  thence  mounted  into  glycerine  jelly. 
Portions  of  the  frond  were  also  embedded  in  pith  and  cut  free- 
hand with  the  razor. 

Some  of  the  material  was  embedded  in  paraffine,  according 
to  the  usual  methods  and  cut  on  the  microtome.  The  sections 
thus  obtained  proved,  in  some  instances,  very  successful,  espe- 
cially in  the  case  of  such  firm  tissues  as  the  stipe  and  the  vege- 
tative portions  of  the  lamina. 

Several  staining  fluids  were  used.  Both  section-staining  and 
staining  in  toto  were  employed.  The  former  method  proved 
most  successful.  The  following  stains  were  found  useful. 

Aniline  blue :  Sections  were  placed  .for  5-10  minutes  in  a 
saturated  solution  of  aniline  blue  (spiritlos)  in  50  per  cent,  alco- 
hol. The  walls  were  stained  a  deep  blue,  gelatinous  structures 
and  cell  contents  a  light  blue.  This  proved  the  most  useful 
stain  for  clearly  defining  anatomical  details. 

Hoffman's  violet :  Sections  were  stained  with  a  saturated  so- 
lution of  Hoffman's  violet  in'  concentrated  sulphuric  acid.  As 
soon  as  the  section  had  taken  a  brown  stain  they  were  placed 
in  water  and  the  acid  washed  out.  The  protoplasm  took  a  blue 
stain,  the  walls  were  unstained.  This  method  proved  useful 
in  staining  the  protoplasmic  connections  between  the  cells. 

Fuchsin :  Dilute  alcoholic  solution  of  fuchsin  stained  the 
walls  light  red,  the  protoplasmic  contents  a  deeper  red. 

Iodine  in  potassium  iodide  :  This  stained  the  floridian  starch 
a  dark  yellow-brown,  the  other  cell  contents  a  light  yellow- 
brown.  It  proved  very  useful  in  staining  the  protoplasmic  con- 
nections between  the  cells,  especially  in  the  gonimoblast  fila- 
ments. 

Gross  anatomy  (Fig.  i). — The  plant  consists  of  a  broad,  flat, 
membranaceous,  more  or  less  subdivided,  blood-red  lamina  borne 
upon  a  short  stipe  which  is  sensibly  continuous  with  the  lamina. 
The  stipe  is  expanded  below  into  a  small  holdfast.  The  lamina 
is  sometimes  nearly  entire  in  outline,  ovate,  or  broadly  lanceo- 
late, sometimes  very  deeply  lobed,  or  divided  almost  to  the  base 
into  2-4  lobes  which  may  be  widely  divergent.  The  lamina  is 
17-40  cm.  long  and  8-17  cm.  wide.  The  two  faces  of  the 
frond  are  in  all  respects  similar.  The  lamina  is  perforated  by 
numerous  holes  ranging  from  .5  mm.  to  10  cm.  in  length  and 


Butters:    OBSERVATIONS  ox  KHODYMENIA.  209 

from  .5  mm.  to  2  cm.  in  width.  The  smallest  are  almost  circu- 
lar in  outline.  Those  somewhat  larger  (up  to  about  5  mm.  in 
diameter)  are  usually  somewhat  oval,  being  elongated  in  the 
direction  of  the  long  axis  of  the  frond.  The  larger  perforations 
are  of  irregular  elongated  outline.  Sometimes  these  perfora- 
tions are  exceedingly  numerous,  as  many  as  four  or  five  per 
square  cm.  being  not  uncommon. 

In  some  specimens  numerous  proliferations  are  borne  on  the 
faces  and  edges  of  the  lamina  and  also  on  the  stipe.  These  are 
cylindrical  bodies,  about  I  mm.  in  diameter,  seldom  more  than 
25  mm.  long,  often  flattened  somewhat  towards  the  apex  and 
often  branched  once  or  twice  or  deeply  lobed  at  the  apex. 

In  two  specimens  the  upper  portions  of  the  laminae  present  a 
peculiar  mottled  appearance  which  was  at  first  supposed  to  be 
due  to  the  presence  of  tetraspores.  It  appears  that  this  is  not 
the  cause  of  the  phenomenon  noted.  This  subject  will  be  dis- 
cussed further  under  the  description  of  the  cystocarp. 

The  entire  stipe  was  present  in  only  one  of  the  specimens  at 
hand.  In  this  it  was  3.5  cm.  long,  about  i  mm.  in  greatest 
diameter,  flattened  somewhat,  parallel  to  the  flat  surface  of  the 
lamina,  passing  insensibly  into  the  lamina  above,  and  expand- 
ing abruptly  below  to  form  the  holdfast,  which  is  a  small,  thin, 
irregular  disk,  about  5  mm.  in  diameter. 

All  of  the  specimens  have  cystocarps  scattered  irregularly 
over  the  entire  surface  of  both  sides  of  the  lamina.  These  are 
bodies  1-1.5  mm.  in  diameter,  nearly  hemispherical,  or  pro- 
truding slightly  at  the  apex,  and  are  extremely  numerous,  as 
many  as  fifteen  being  often  found  on  one  square  cm.  of  the 
lamina. 

Minute  anatomy :  Lamina. — The  lamina  consists  of  pseudo- 
parenchymatous  tissue  of  which  two  principal  areas  may  be  dis- 
tinguished in  the  cross-section  (a),  a  central  layer  of  large- 
celled  ;  and  (b)  a  cortical  layer  of  small-celled  tissue  (Fig.  2). 

(a)  The  cells  of  the  central  area  are  large,  generally  some- 
what flattened  parallel  to  the  surface  of  the  frond,  isodiametrical 
in  tangential  section  (Fig.  3).  The  cells  vary  greatly  in  size, 
the  average  being  73.5  x  105  mic.,  while  cells  occur  as  small  as 
6x  14  mic.,  and  as  large  as  100  x  200  mic.  The  largest  cells 
are  situated  near  the  central  portion  of  this  area,  and  from  them 
the  size  of  the  cells  decreases  quite  regularly  towards  the  more 
superficial  portions.  The  more  superficial  cells  of  this  area 


210  MINNESOTA    BOTANICAL    STUDIES. 

differ  considerably  in  other  respects  also,  from  the  central  cells. 
The  outer  cells  are  more  flattened  than  the  central  ones,  they 
are  more  densely  protoplasmic  and  are  filled  with  grains  of 
floridian  starch,  while  but  little  starch  occurs  in  the  central  cells. 
In  this  more  superficial  portion  of  the  central  area  the  proto- 
plasmic connections  between  the  cells  of  the  filaments  of  which 
the  tissue  is  composed  can  be  easily  made  out.  They  are 
plainly  visible  in  both  stained  and  unstained  preparations,  and 
in  both  tangential  and  cross  sections.  In  accordance  with  the 
less  dense  protoplasmic  contents  of  the  central  cells,  the  con- 
necting strands  are  less  evident  among  them,  but  when  the  walls 
of  this  area  are  stained,  numerous  pits  are  shown  penetrating 
the  walls  of  the  central  cells  (Fig.  3).  These  pits  appear  to  be 
of  irregular  distribution  and  often  more  than  one  are  to  be  ob- 
served between  the  same  pair  of  cells.  They  can  be  best  ob- 
served in  a  tangential  section. 

(b)  The  cortical  area  consists  of  small  cells  almost  spherical 
or  with  the  longest  diameter  perpendicular  to  the  surface  of  the 
frond,  of  quite  uniform  size  (averaging  5.7  x8-5  mic.),  arranged 
in  1-3  layers,  either  in  filaments  perpendicular  to  the  surface  of 
the  frond  or  somewhat  irregularly.  In  a  surface  view  of  the 
lamina  they  appear  entirely  irregular  in  arrangement  (Fig.  4). 
The  cells  are  densely  protoplasmic  and  contain  chromatophores. 

Stipe. — The  general  structure  of  the  stipe  (Fig.  5)  is  similar 
to  that  of  the  lamina,  but  there  are  numerous  special  modifications 
of  the  several  areas.  The  cells  of  the  central  area  are  elongated 
somewhat  in  the  direction  of  the  axis  of  the  frond  and  are  some- 
what compressed  parallel  to  the  compression  of  the  stipe. 
They  are  of  more  uniform  size  than  were  those  of  the  lamina, 
and  they  were  otherwise  more  nearly  uniform  than  those  of  the 
lamina.  Their  average  size  is  57  x86  x  143  mic.  The  cortical 
area  of  the  stipe  is  much  thicker  than  that  of  the  lamina.  It  is  3-8 
cells  deep.  The  cells  are  larger  than  the  corresponding  cells 
of  the  lamina  (average  14x23  mic.)  and  are  conspicuously  ar- 
ranged in  filaments  running  perpendicular  to  the  surface  of  the 
stipe. 

Proliferations. — The  structure  of  the  proliferation  is  similar 
to  that  of  the  main  frond.  The  central  cells  are  somewhat 
elongated  in  the  direction  of  the  axis,  and,  in  general,  are  more 
numerous  and  smaller  than  the  corresponding  ones  of  the  main 
frond.  All  the  cells  except  those  of  the  cortical  area  contain 
considerable  floridian  starch. 


Butters:   OBSERVATIONS  ON  RHODYMENIA.  211 

Reproductive  tract. — Cystocarp  (Fig.  7-12).  The  cystocarp 
projects  on  one  side  of  the  thallus.  The  pericarp  is  composed 
of  thickened  cortical  tissue,  which,  in  the  mature  cystocarp,  is 
8-40  cells  thick.  The  outer  walls  of  the  pericarp  are  small 
and  resemble  those  of  the  cortex  of  the  vegetative  part  of  the 
frond.  The  inner  cells  are  large  (14  x  29  mic.),  flattened  paral- 
lel to  the  surface  of  the  cystocarp  and  show  numerous  irregu- 
lar protoplasmic  connections.  The  cystocarp  opens  by  a  carpo- 
stome  situated  at  the  apex  of  the  pericarp. 

The  sporogenous  tissue  is  in  the  form  of  an  irregularly  lobed 
mass,  borne  on  a  basal  placenta,  and  partially  filling  the  cavity 
of  the  cystocarp.  The  space  between  the  spore  mass  and  the 
pericarp  is  filled  with  gelatine.  The  placenta  is  a  mass  of  small- 
celled  tissue  containing  numerous  intercellular  spaces,  which 
rests  upon  the  large-celled  central  tissue  of  the  lamina.  The 
gonimoblast  filaments  (Fig.  9-11)  branch  repeatedly.  They 
consist  of  irregular  elongated  or  rounded,  often  club-shaped 
cells,  with  dense,  finely  granular  protoplasm  and  very  trans- 
parent walls.  They  contain  no  starch.  They  are  connected 
in  filaments  by  very  broad  protoplasmic  connections  surrounded 
midway  by  a  callous-like  ring.  The  upper  cells  of  the  fila- 
ments are  smaller  and  more  rounded  than  the  lower.  The 
structure  of  the  filaments  was  best  shown  in  preparations  made 
by  pressing  out  the  contents  of  a  mature  cystocarp  upon  a  mi- 
croscopic slide,  staining  lightly  with  iodine  in  potassium 
iodide,  and  then  pressing  out  with  a  cover  glass.  The  spores 
themselves  are  irregular,  ovoid,  thin-walled  cells,  densely 
packed  with  floridian  starch.  Their  average  size  is  21.5  x  34.5 
mic.,  but  they  vary  considerably  in  this  respect.  The  number 
of  spores  produced  in  each  cystocarp  is  very  great,  20,000-30,- 
ooo  being  not  uncommon.  The  lowest  lobes  of  the  sporogenous 
mass  appear  to  be  in  all  cases  sterile.  They  form  small  masses 
of  compact  tissue  consisting  for  the  greater  part  of  cells  having 
about  one-half  the  diameter  of  mature  spores  and  containing 
but  little  starch.  A  few  long  cells  like  those  of  the  goni- 
moblast filaments  also  occur  in  this  region. 

In  many  cystocarps  branched  filaments  of  cells  rise  from  the 
vegetative  tissue  at  the  base  of  the  cystocarp.  The  cells  of 
these  filaments  (Fig.  8)  exhibit  peculiar  lateral  outgrowths 
which  appear  to  fuse  with  the  adjacent  cells  or  with  similar 
outgrowths  from  them,  forming  a  peculiar  loose,  irregularly 


212  MINNESOTA    BOTANICAL    STUDIES. 

connected  tissue  similar  to  that  which  forms  the  inner  portion  of 
the  cystocarp  wall.  These  filaments  are  not  present  in  all  cys- 
tocarps.  They  appear  to  result  from  the  tearing  of  the  sub- 
cortical  tissue  in  the  formation  of  the  cavity  of  the  cystocarp. 

In  the  young  cystocarp  (Fig.  12)  the  thickened  wall  is  al- 
ready present  and  shows  its  permanent  division  into  two  layers. 
The  cells  of  the  outer  layer  are  arranged  in  filaments  perpen- 
dicular to  the  surface,  those  of  the  subcortical  layer  are  ar- 
ranged in  oblique  rows  converging  towards  the  apex  of  the 
pericarp.  The  placental  area  and  spore  mass  are  represented 
by  a  few  connected  cells  with  very  dense  contents.  The  cavity 
of  the  cystocarp  is,  at  this  stage,  comparatively  small.  The 
carpostome  is  already  developed  even  in  very  young  cystocarps. 
It  appears  to  be  formed  by  the  tearing  apart  of  the  cells  to- 
gether with  the  destruction  of  some  of  the  cells.  As  was  men- 
tioned in  the  account  of  the  gross  anatomy  of  the  lamina,  in 
two  specimens  the  apical  portions  of  the  frond  present  a  pecu- 
liar mottled  appearance.  Cross  sections  of  these  areas  show 
that  in  places  there  are  slight  protuberances  from  the  surface  of 
the  thallus  associated  with  an  unusual  development  of  cortical 
and,  in  some  cases,  also  of  the  subcortical  cells.  The  cortical 
cells  are  rather  narrower  than  elsewhere,  and  more  elongated 
perpendicularly  to  the  surface  of  the  frond.  The  cortical  layer 
is  also  a  greater  number  of  cells  deep  than  elsewhere,  and  in 
some  cases  there  is  also  increase  in  the  number  of  the  smaller 
central  cells  situated  immediately  beneath  the  cortex.  These 
areas  in  some  cases  involve  only  a  few  cells,  in  others  they  are 
.5  mm.  in  diameter.  The  structure  of  the  larger  protuberances 
agrees  essentially  with  that  of  the  young  cystocarp  described 
above,  except  that  in  no  cases  could  any  cavity  be  discovered 
in  them.  They  appear  to  me  to  be  very  young  stages  of  cysto- 
carp development,  but  in  no  case  could  positive  evidence  of 
their  nature  be  discovered. 

No  indubitable  cases  of  tetraspore  formation  were  seen,  but 
in  some  cross  sections  taken  through  the  upper  part  of  the 
lamina,  some  of  the  cortical  cells  are  peculiarly  divided  pro- 
ducing somewhat  the  appearance  of  tetraspore  formation  (Fig. 
6).  Except  for  their  peculiar  arrangement,  these  cells  appear 
in  all  respects  similar  to  the  ordinary  cortical  cells.  Whether 
they  are  tetraspores  or  not,  could  not  be  determined.  Their 
method  of  division  is  cruciate  or  somewhat  irregular  approach- 
ing the  tetrahedral  arrangement. 


Butters:     OBSERVATIONS    ON    RHODYMENIA.  213 

BIBLIOGRAPHY. 

Greville,  R.  K.     Alg.      1830. 

Montague,  C.     Crypt.   Brasil.  in  Ann.  Sci.  Nat.  Bot.  II.  12:  42- 

55-      lS39- 

Postels  A.  and  Ruprecht,  F.   J.     Illust.  Algar.  20.  pf.  j6.     1840. 
Kiitzing,  F.  T.     Spec.  Algar.  693,  778-784.      1849. 

(1)  Agardh,  J.  G.     Alg.  Med.  et  Adriat.      1842. 

(2)  Agardh,  J.  G.     Alg.  Liebm. 

(3)  Agardh,  J.  G.     Spec.  Gen.  et  Ord.  Algar.  2:  374-383.     1852. 

(4)  Agardh,  J.  G.     Epicr.  syst.  Florid.  324,  329.      1876. 
Harvey,  W.  H.     Ner.  Bor.-Am.  2:    147,  148.     1853. 

Harvey,  W.  H.  Coll.  Algae  Northwest  Coast  of  North  America, 
in  Journ.  Linn.  Soc.  Bot.  6:  157-177.  1862. 

Kjellman,  F.  R.     Alg.  Arctic  Sea.      150,  151.      1883. 

Ardissone,  F.,  Phyc.  Med.  i:   212-215.      I^3- 

De-Toni,  J.  B.  and  Levi,  D.  Schem.  Gen.  Florid.  Illustr.  Nota- 
risia.  3:  appendix  I-XXV.  1888. 

Carruthers,  J.  B.  On  the  cystocarps  of  species  of  Callophyllis  and 
Rhodymenia.  Jour.  Linn.  Soc.  Bot.  29:  77-86.  1893. 

Osterhout,  W.  J.  V.  A  simple  freezing  device.  Bot.  Gaz.  21 : 
195-201.  1896. 

DESCRIPTION  OF  PLATE  XX. 

1.  Mature  plant    showing  perforations,   proliferations    and  cysto- 
carps.    x  \. 

2.  Cross   section  of  frond.      Med.    medullary   area,    cor.    cortical 
area,     x  1 12. 

3.  Longitudinal  section  of  central  area  of  frond,  showing  pits  in  the 
walls,     x  132. 

5.  Surface  view  of  frond,  showing  irregular  arrangement  of  surface 
cells.  Stained  with  fuchsin.  X335. 

5.  Cross  section  of  stipe.     Drawn  with  camera  lucida.     X4O. 

6.  Tetraspores  ?     X335. 

7.  Cross  section  of  mature  cystocarp  (not  cut  through  carpostome) 
showing  placentation  and  general  structure  of  the  spore  mass.      The 
upper  part  of  the  spore  mass  is  somewhat  scattered.     Only  the  spores 
have  their  contents  filled  in.     X39« 

8.  Peculiarly  branched  and  interwoven  filaments  from  the  base  of  a 
cystocarp.     x  237. 

9.  Gonimoblast  filaments  stained  one  minute  in  iodine  in  potassium 
iodide,      x  237. 

10.  ii.   Gonimoblast  filaments,  contents  not  drawn.     X237. 

12.  Young  cystocarp  showing  carpostome  and  early  stage  in  the 
development  of  spore  mass.  Contents  have  been  omitted  except  from 
sporogenous  cells.  Di'awn  with  Camera  lucida.  x  230. 


VOL  11 


MINNESOTA 


aroqi 


AN1CAL  STUDIES. 


PART  III. 


XX. 


NO   CO..    BOSTON. 


XVIII.     CONTRIBUTIONS    TO    A    KNOWLEDGE    OF 

THE  LICHENS   OF   MINNESOTA.— IV.    LICHENS 

OF   THE   LAKE   SUPERIOR   REGION. 


BRUCE  FINK. 


CONSIDERATIONS   OF  DISTRIBUTION  AND   HABITAT. 

The  area  treated  in  this  paper  includes  essentially  the  counties 
of  Cook  and  Lake,  comprising  about  5,000  square  miles  of  land. 
It  lies  to  the  northwest  of  lake  Superior,  bordering  on  the  lake 
for  about  150  miles  and  on  the  province  of  Ontario,  Canada,  for 
about  125  miles,  thus  forming  the  extreme  northeastern  portion 
of  the  State  of  Minnesota.  It  was  supposed,  before  studying 
it,  that  the  region,  because  of  its  position,  would  furnish  many 
lichen  species  new  to  the  State  and  to  the  interior  of  North 
America,  and  the  investigation  has  fulfilled  expectations. 
Besides  its  geographical  position,  certain  physical  features 
have  produced  diversity  of  lichen  species  as  will  be  shown  later. 

The  lichens  of  the  region  have  never  been  studied  previously. 
Tuckerman,  in  his  Synopsis,*  mentions  collections  from  the 
north  shore  of  lake  Superior  by  John  Macoun  and  L.  Agassiz. 
The  collections  by  Agassiz  were  made  in  1848  and  published  f 
from  1850  to  1852.  He  traversed  the  north  shore  from  Sault 
Saint  Marie  to  Fort  William.  The  collecting  by  Macoun  was 
done  in  July,  1869,  along  the  north  shore  in  Canada,  and  in 
1884  around  lake  Nipigon.  A  publication  may  soon  be  ex- 
pected from  the  latter  collector,  giving  a  complete  list  of  the 
Canadian  lichens,  and  this  should  add  much  of  interest  con- 
cerning some  species  listed  in  this  paper.  Of  the  two  collec- 
tors, Agassiz,  at  Fort  William,  came  within  about  50  miles, 
while  Macoun  probably  came  within  150  miles  of  certain  points 


*  Tuckerman,  E.,  Synopsis  of  the  North  American  Lichens,  Parts  I  and  II, 
1882  and  1888. 

f  Agassiz,  L.,  Lake  Superior,  its  Physical  Character,  Vegetation  and  Animals 
compared  with  those  of  other  and  similar  Regions,  pp.  170-174.  Boston,  1850. 

215 


216  MINNESOTA    BOTANICAL    STUDIES. 

reached  by  me  along  the  international  boundary  or  the  shore  of 
the  lake. 

Several  species  of  the  lichens  collected  by  Dr.  C.  C.  Parry 
in  1848  and  published  in  1895  *  show  northern  range,  but  care- 
ful investigation  f  indicates  that  they  were  collected  south  of 
lake  Superior.  Thus  it  appears  that  the  collections  listed 
herein  are  the  first  made  on  the  north  shore  of  lake  Superior 
in  Minnesota,  or  along  the  international  boundary  for  more  than 
100  miles  west  of  the  lake. 

The  collecting  was  done  by  the  writer,  assisted  by  Mr.  A.  S. 
Skinner,  during  the  latter  part  of  June  and  the  whole  month  of 
July,  1897.  We  were  fortunate  enough  to  be  associated  with 
Dr.  A.  H.  Elftman,  who  wished  to  traverse  the  region  for  geo- 
logical study  and  whose  thorough  knowledge  of  the  territory 
covered,  alone  made  it  possible  for  us  to  find  the  best  collecting 
stations  in  this  for  the  most  part  uninhabited  region,  and  thus  to 
accomplish  good  results  in  a  comparatively  short  time. 

We  reached  Grand  Portage  island  on  the  north  shore  of  lake 
Superior,  June  i6th,  and  began  collecting  at  once.  The  plan 
was  to  study  the  lichen  flora  of  the  international  boundary  and 
the  north  shore  of  lake  Superior  within  the  boundaries  of 
Minnesota  and  to  reach  some  of  the  inland  portions  of  the  two 
counties.  On  the  whole  trip  we  sought  localities  as  collecting 
stations  offering  the  greatest  differences  as  to  elevation,  rock 
formations,  arboreal  flora,  soil,  moisture,  etc.  Beginning  at 
Grand  Portage  island  we  traveled  westward  by  Pigeon  river  and 
the  chain  of  lakes  along  the  international  boundary  to  Gunflint, 
thence  south  by  a  series  of  lakes  to  Poplar  river  and  down  the 
river  to  Lutsen,  on  the  shore  of  lake  Superior.  From  here  we 
proceeded  down  the  lake  to  Tofte,  Beaver  Bay  and  Two  Har- 
bors. We  next  went  directly  to  Ely  and  thence  eastward  to 
Snowbank  lake.  We  made  as  thorough  a  study  as  possible  of 
the  lichen  flora  of  the  Grand  Portage  area,  and  then  stopped  for 
study  whenever  and  wherever  we  found  enough  of  difference  in 


*Fink,  B.,  Lichens  collected  by  Dr.  C.  C.  Parry  in  Wisconsin  and  Minne- 
sota in  1848.  Proc.  Iowa  Acad.  Sci.  2  :  137.  1895. 

t  Parry,  C.  C.,  Systematic  Catalogue  of  Plants  of  Wisconsin  aud  Minnesota, 
made  in  connection  with  the  geological  survey  of  the  Northwest  during  the 
season  of  1848.  In  Owen,  D.  D.,  Report  of  a  geological  survey  of  Wisconsin, 
Iowa,  Minnesota  and  incidentally  of  a  portion  of  Nebraska  Territory.  Appen- 
dix, article  V.  606-622.  1852,  mentions  Cladonia  rangiferina  (L. )  Hoffm- 
and  Gyrophora  muhlenbergii  Ach.  from  Falls  of  St.  Croix. 


Fink  :     LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  217 

physical  environment  to  lead  to  the  conclusion  that  time  would 
be  profitably  employed. 

The  writer  was  relieved  of  camp  duty  as  much  as  possible, 
so  that  there  was  some  time  for  collecting  each  day,  even  when 
traveling.  However,  the  few  collections  thus  made  are  nearly 
all  recorded  in  the  list  with  those  of  the  nearest  well  studied 
locality.  The  principal  collecting  stations  are  given  below,  with 
elevation  and  time  spent  in  collecting  for  each.  Since  many  of 
these  stations  are  in  uninhabited  and  little  known  regions,  I  have 
given  the  township  and  range  of  each  one. 

I.  Grand  Portage  and  Grand  Portage  island,  9  days,  elevation 

602  to  1305  feet,  T.  63  N.,  R.  6  E. 

II.  English  portage,  3  hours,  elevation  1339  feet,  T.  64  N.,  R. 

4E. 

III.  South   Fowl  lake,  3  hours,  elevation  1436-1450  feet,  T. 

64  N.,  R.  4  E. 

IV.  Moose  lake,   2  hours,  elevation    1492   feet,  T.  65  N.,  R. 

3E. 

V.  Rose  lake,  i  day,  elevation   1528  feet,  T.  65  N.,  R.  i  W. 

VI.  Paulson  iron  mines,  2  days,  elevation  1825  to  2000  feet, 

T.  65  N.,  R.  4  W. 

VII.  Gunflint,  i  day,  elevation  1547  to  1650  feet,  T.  65  N.,  R. 

3  W. 

VIII.  Winchell   lake,  2   days,  elevation  1910  to  2230  feet,  T. 

64  N.,  R.  2  W. 

IX.  Brule  lake,  5   hours,   elevation   2084  ^eet'  T.  63   N.,  R. 

3  W. 

X.  Tofte,  3  days,  elevation  927  to  1529  feet,  T.  59  N.,  R.  4  W. 

XI.  Beaver  Bay,  2  days,  elevation  602  to  1250  feet,  T.  55  N., 

R.  8  W. 

XII.  Great  Palisades,  6  hours,  elevation  602  to  1200  feet,  T.  56 

N.,  R.  7  W. 

XIII.  Two  Harbors,  2  hours,  elevation  692  feet,  T.  52  N.,  R. 

ii  W. 

XIV.  Prairie  portage,  i  day,  elevation  1300  feet,  T.  64  N.,  R. 

9  W. 

XV.  Iron  Mountain  lake,  i  day,  elevation  1342  feet,  R.  64  N., 

R.  8  W. 

XVI.  Snowbank  lake,  4  days,  elevation  1424  feet,  T.  64  N., 
R.  9  W. 

XVII.  Disappointment  lake,  i   day,  elevation  1449-1850  feet, 
T.  64N.,R.  8  W. 


218  MINNESOTA    BOTANICAL    STUDIES. 

XVIII.  Moose  lake,  i  day,  elevation,  1339  feet,  T.  64  N.,  R. 
9  W. 

XIX.  Wind  lake,   i    day,  elevation  1359   feet,  T.   64  N.,  R. 
9  W. 

XX.  Ella   Hall  lake,  3  hours,  elevation  1306  feet,  T.  64  N., 

R.  10  W. 

XXI.  Fall  lake,  3  hours,   elevation   1313   feet,  T.  63  N.,  R. 

ii  W. 

Of  the  stations  given  above,  numbers  XIV  to  XIX  inclusive 
have  been  designated  in  the  list  of  species  as  the  Snowbank 
lake  area,  XX  and  XXI  as  Ely,  VIII  and  IX  as  the  Misquah 
hills,  and  VI  and  VII  as  Gunflint.  All  other  areas  include 
each  but  a  single  station.  The  quantity-collecting  being  largely 
done  when  we  left  Grand  Portage,  we  were  able  to  move  rapidly, 
as  only  plants  not  found  in  this  first  area  needed  to  be  collected 
in  bulk.  For  illustration  of  distribution,  the  collections  were 
made  as  complete  as  possible  at  Grand  Portage,  Gunflint,  in  the 
Misquah  hills,  at  Tofte,  at  Beaver  Bay  and  in  the  Snowbank 
lake  area. 

There  is  an  appreciable  difference  between  the  lichen  flora  of 
Grand  Portage  island  and  that  of  the  mainland  two  miles  across 
the  bay.  The  island  reaches  an  elevation  of  only  125  feet  above 
lake  Superior  while  Mt.  Josephine  on  the  mainland  reaches  an 
altitude  of  about  800  feet  above  the  lake.  The  Keweenawan 
series  of  rocks,  which  appears  on  the  island,  is  wanting  on  the 
portion  of  the  adjacent  mainland  explored,  while  the  Animikie 
series  is  found  in  both  places.  However,  I  could  not  ascertain 
that  difference  in  petrographical  construction  in  any  noticeable 
way  determines  the  floral  differences  either  here  or  elsewhere  in 
the  territory  studied.  Passing  by  the  Gunflint  area  for  the  pres- 
ent, I  may  say  that  the  Misquah  hills  were  regarded  as  especi- 
ally important,  since  they  contain  the  highest  areas  in  the  state, 
and  were  as  carefully  studied  as  our  time  would  permit.  Carl- 
ton  peak  at  Tofte,  and  the  Palisades  were  points  of  special  in- 
terest. Two  Harbors  was  of  interest  as  it  is  the  most  southern 
point  reached  in  the  survey,  and  Ely  was,  also,  as  it  is  the  most 
western.  However,  the  Snowbank  lake  area  somewhat  further 
east  was  much  more  thoroughly  studied  than  Ely. 

On  the  whole  all  of  the  two  counties  was  studied  thoroughly 
enough  to  know  that  practically  all  of  the  lichens  generally  dis- 
tributed over  the  area  were  secured  as  well  as  many  more  which 


Fink  :     LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  219 

as  yet  show  only  local  occurrence.  Examination  of  the  route 
will  show  that  we  covered  all  of  the  international  boundary  be- 
tween Ely  and  Grand  Portage,  except  about  20  miles  in  a 
straight  line  from  the  most  eastern  portion  of  the  Snowbank 
lake  area  to  Gunflint.  Thus  the  boundary  was  well  studied. 
The  line  of  travel  through  the  Misquah  hills  from  Gunflint  to 
Lutsen  gave  a  fair  view  of  the  interior  of  the  region  as  well  as 
the  highest  area  in  the  state.  A  day  .spent  along  the  shore  at 
Lutsen  failed  to  furnish  any  species  not  found  at  Grand  Port- 
age. Consequently,  as  we  were  to  stop  at  Tofte,  only  10  miles 
distant,  no  record  was  made  of  the  species  found  at  the  former 
place.  Tofte,  Beaver  Bay  and  the  Palisades  gave  a  good  view  of 
the  lake  shore  and  higher  elevations  near  by  in  the  Sawteeth 
mountains.  It  is  to  be  regretted  that  we  did  not  have  oppor- 
tunity for  examination  of  the  lake  shore  and  Sawteeth  moun- 
tains at  some  points  between  Lutsen  and  Grand  Portage,  but 
doubtless  the  number  of  additional  species  would  not  have  been 
large  after  a  thorough  examination  of  the  shore  both  to  the  ex- 
treme north  and  toward  the  south  of  the  area  studied. 

The  whole  region  is  one  of  extreme  interest  to  the  lichenist 
because  of  the  diversity  of  natural  conditions  which  gives  a  flora 
rich  in  individuals  as  well  as  variations  within  certain  species 
which  attracted  special  attention.  The  great  masses  of  igneous 
and  metamorphic  rocks  along  the  Superior  and  inland  shore 
lines,  the  same  rocks  back  from  shore  lines  and  the  coniferous 
and  various  other  trees  together  with  diversity  as  to  temperature, 
moisture  and  elevation,  all  help  to  produce  a  flora  richer  in 
lichen  species  than  I  had  expected  to  find.  Though  the  annual 
precipitation  of  moisture  for  the  area  is  not  large,  yet  the  com- 
paratively impervious  nature  of  the  rocks  causes  the  water  to 
collect  in  depressions  of  surface,  forming  a  multitude  of  lakes 
of  various  sizes  whose  moist  borders  are  a  veritable  paradise  for 
lichens  and  especially  for  lithophytic  species.  The  dense 
forests  also  hold  moisture  and  favor  lichen  growth.  When  one 
finds  single  branches  of  Usnea  longissima  Ach.  five  feet  long, 
as  we  collected  on  Grand  Portage  island,  he  realizes  the  signifi- 
cance of  the  name.  Here  and  in  some  other  localities  of  the 
region  studied  the  dying  conifers  especially  are  literally  covered 
with  this  plant,  other  species  of  the  genus  and  Alectoria  jubata 
(L.)  Nyl.,  all  growing  in  a  tangled  profusion  which  obscures 
the  host  and  when  wet  with  rain  or  dew  furnishes  a  view  of  sur- 


220  M  NNESOTA    BOTANICAL    STUDIES. 

passing  beauty.  Hardly  less  remarkable  is  the  growth  of  Cla- 
donia rangiferina  (L.)  Hoffm.  in  open  woods  near  Mt.  Josephine, 
single  clusters  measuring  three  or  four  feet  across  and  reaching 
a  foot  in  height.  This  plant  was  also  common  on  rocks  and  in 
crevices  exposed  to  wind  and  sun,  but  was  always  much  smaller 
in  such  locations.  It  is  evidently  not  a  natural  pioneer  among 
lichens,  but  grows  after  other  plants  have  attacked  the  rocky 
substratum,  or  on  a  thin  layer  of  soil  in  crevices,  and  best  of  all 
after  trees  or  shrubs  have  grown  sufficiently  to  protect  it  some- 
what from  wind  and  sun  and  have  not  yet  become  large  enough 
or  thick  enough  to  kill  it  out.  This  same  kind  of  ecological 
relation  favors  Cladoniafurcata  (Huds.)  Fr.,  a  variety  of  which 
was  found  fruiting  only  in  such  environment.  More  is  given 
below  about  other  Cladonias,  and  the  observation  could  be 
extended  to  Stereocaulon. 

After  fires  have  passed  over  a  region  destroying  the  trees  and 
small  scattered  second  growth  begins  to  appear  to  furnish  some 
protection,  Cladonia  cristatella  Tuck,  and  a  large  variety  of 
forms  of  C.  gracilis  (JL.)  Nyl.  soon  begin  to  grow  in  great  pro- 
fusion on  old  stumps,  prostrate  logs  and  bits  of  decaying  wood 
lying  upon,  or  more  or  less  sunken  into  the  soil.  Only  a  few  of 
the  many  varieties  of  the  latter  plant  allowed  by  European 
lichenists  are  recognized  in  the  list  of  species  though  forms 
closely  resembling  other  varieties,  so  called,  are  represented  in 
my  collections.  Nothing  seems  to  be  gained  by  carrying  the 
"  splitting"  process  to  extremes  without  a  study  of  life  histor- 
ies. 6\  gracilis  (L.)  Nyl.  in  regions  recently  burned  showed 
much  less  variation  than  in  places  where  the  species  had  been 
established  longer  since  the  burning,  and  a  careful  study  of  a 
large  number  of  individuals  in  this  region,  extending  over  a  ser- 
ies of  years  would  enable  one  to  trace  the  growth  and  variation 
within  single  individuals  and  thus  establish  varieties  with  cer- 
tainty. Great  variety  was  observed  in  the  plants  in  regions 
that  had  burned  15  or  at  most  20  years  ago  so  that  a  study  ex- 
tending over  10  years  should  be  sufficient  to  give  the  desired 
data. 

Like  Cladonia  rangiferina  (L.)  Hoffm.,  C.  cristatellaTvick.. 
is  extremely  sensitive  to  environment.  In  regions  where  the 
plants  are  exposed  to  sun  and  wind  and  in  stations  of  high  ele- 
vation, the  plants  are  much  smaller  than  in  better  shaded  and 
less  elevated  places.  The  relation  of  size  to  amount  of  protec- 


Fink  :     LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  221 

tion  offered  by  trees  and  shrubs  may  be  observed  in  many  places 
between  Gunflint  and  Lutsen,  especially  at  Gunflint  and  in  the 
Misquah  hills.  The  effect  of  elevation,  or  rather  the  combined 
effect  of  elevation  and  exposure,  was  especially  noted  on  Mt. 
Josephine  and  on  Carlton  peak.  In  both  of  these  places  the 
stunted  condition  was  also  noticeable  in  other  Cladonias  and  in 
lichens  belonging  to  other  genera. 

The  part  that  lichens  play  in  rock  decay  and  soil  formation 
was  studied  in  a  general  way  in  the  Grand  Portage  area,  and 
some  of  the  most  noticeable  facts  are  stated  below.  Grand 
Portage  island  contains  57  acres  of  land  and  furnishes  sufficient 
variety  as  to  substrata  suitable  to  lichen  growth  to  make  the  study 
interesting.  The  crustaceous  lichens  furnish  most  of  the  species 
which  first  gain  a  footing  on  the  rocks,  and  of  these  were  found 
on  the  island  three  or  four  species  of  Placodium,  a  half  dozen 
or  more  rock  Lecanoras  as  well  as  a  larger  number  of  Biatoras, 
Lecideas  and  Buellias.  Of  the  foliaceous  lichens  the  Umbili- 
carias  are  most  characteristically  rock  pioneers ;  but  these  were 
very  rare  on  the  island,  which  did  not  furnish  the  high  bluffs 
that  they  seek  especially.  As  soon  as  rock  decay  has  begun, 
the  less  strictly  crustaceous  species  begin  to  appear.  Of  these 
Pannaria  microphylla  (Sw.)  Delis  was  especially  noticed  some- 
times growing  on  quite  firm  rock,  but  more  frequently  on  rotten 
rock  or  a  residual  product  of  rock  decay  still  in  situ  and  pro- 
tected by  the  lichen  though  sometimes  several  inches  deep. 
Next  come  the  typically  foliaceous  and  fruticulose  species  as  the 
Peltigeras  and  Cladonias.  Finally  enough  soil  is  established 
so  that  smaller  Spermaphytes  and  finally  trees  and  shrubs  be- 
come established,  these  larger  ones  in  turn  furnishing  substrata 
for  epiphytic  lichens.  At  the  present  time,  trees  grow  at  one 
end  of  the  ridge  of  highest  land  extending  across  the  island 
while  the  other  end  is  bare  of  trees  and  soil  to  a  large  extent 
and  yet  supports  many  strictly  lithophytic  lichens.  At  the  shore 
line  one  finds  amphibious  Endocarpons  and  a  Collema  while 
typically  xerophytic  species  cover  the  remainder  of  the  island. 
The  analysis  could  be  extended  to  include  a  statement  of  differ- 
ent sorts  of  woody  substrata  which  result  in  giving  diversity  of 
lichens  growing  on  wood  and,  indeed,  to  give  a  detailed  account 
of  substrata  including  that  of  each  one  of  the  88  species  and 
varieties  listed  from  the  island.  But  this  would  lead  to  more 
detail  than  can  be  undertaken  here,  and  for  more  minute  ac- 


222  MINNESOTA    BOTANICAL    STUDIES. 

count  I  have  been  compelled  to  select  very  small  islands,  though 
not  offering  so  much  diversity  as  to  substrata,  and  have  even 
then  confined  the  analysis  to  the  lithophytic  and  a  few  epigean 
species. 

For  this  study  of  islands  three  were  selected  in  the  Snowbank 
lake  area,  and  the  lithophytic  species  were  carefully  noted  on 
two  of  them  and  on  the  other  also  the  decrease  in  number  due 
to  the  establishment  of  an  arboreal  flora.  It  is  to  be  regretted 
that  the  study  could  not  have  been  extended  to  more  islands  and 
to  include  epiphytic  and  epigean  species  as  well  as  lithophytic. 
Island  number  one  is  situated  in  Sucker  lake,  30  feet  from  the 
shore,  in  the  N.  W.  ^  of  S.  W  ^  of  S.  E.  ^  of  Sec.  i,  T.  64 
N.,  R.  9  W.  The  size  of  the  island  is  about  70x75  feet.  The 
surface  is  rocky  with  soil  in  a  few  places  formed  in  situ  or 
washed  in  from  the  lake,  so  that  Cladonias  were  well  estab- 
lished. About  twenty  shrubs  were  growing  on  the  island  and 
two  rather  small  pines.  The  species  noted  in  a  short  time  are 
as  follows  : 

1.  Cladonia  rangiferina  (L.)  HOFFM. 

2.  Cladonia  rangiferina  (L.)  HOFFM.  var.  sylvatica  L. 

3.  Cladonia  rangiferina  (L.)  HOFFM.  var.  alpestris  L. 

4.  Cladonia  pyxidata  (L.)  FR. 

5.  Cladonia  gracilis  (L.)  FR. 

6.  Cladonia  uncialis  (L.)  FR. 

7.  Stereocaulon  paschale  (L.)  FR. 

8.  Umbilicaria  muhlenbergii  (Acn.)  TUCK. 

9.  Endocarpon  fluviatile  DC. 

10.  Parmelia  conspersa  (EHRH.)  ACH. 
n.  Parmelia  saxatilis  (L.)  FR. 

12.  Parmelia  caperata  (L.)  ACH. 

13.  Physcia  sp. 

14.  Physcia  stellaris  (L.)  TUCK. 

15.  Physcia  speciosa  (WuLF.,  ACH.)  NYL. 

16.  Physcia  obscura  (EHRH.)  NYL. 

17.  Ephebe  solida  BORN. 

18.  Pannaria  microphylla  (Sw.)  DELIS. 


Fink :   LICHENS  OF  THE  LAKE  SUPERIOR  REGION.  223 

19.  Urceolaria  scruposa  (L.)  NYL. 

20.  Placodium  vitellinum  (EHRH.)  NAEG.  and  HEPP. 

21.  Lecanora  rubina  (ViLL.)  ACH. 

22.  Lecanora  cinerea  (L. )  SOM.MERF. 

23.  Buellia  petraea  (FLOT.,  KOERB.)  TUCK. 

Island  number  two  is  in  Snowbank  lake,  50  feet  from  the 
shore,  in  the  N.  W.  ^  N.  E.  ^  of  Sec.  29,  T.  64  N.,  R.  8  W. 
near  the  outlet  of  the  lake.  The  size  of  the  island  is  about 
80  x  100  feet,  and  it  is  thickly  covered  with  trees  and  shrubs 
except  in  a  few  spots  where  Cladonia  rangiferina  (L.)  Hoffm. 
persists.  The  species  listed  below  for  this  island  are  excepting 
the  Cladonia )  confined  to  a  circle  of  rock  extending  around  the 
island  and  up  from  the  water  three  inches  to  one  foot.  The 
species  are  as  follows  : 

1.  Cladonia  rangiferina  (L.)  HOFFM. 

2.  Endocarpon  fluviatile  DC. 

3.  Parmelia  conspersa  (EHRH.)  ACH. 

4.  Parmelia  caperata  (L.)  ACH. 

5.  Physcia  obscura  (EHRH.)  NYL. 

6.  Leptogium  lacerum  (Sw.)  Fr. 

7.  Placodium  aurantiacum  (LIGHT.)  NAEG.  and  HEPP. 

8.  Lecanora  subfusca  (L.)  ACH. 

9.  Lecanora  cinerea  (L.)  SOMMERF. 

Island  number  three  is  in  Disappointment  lake,  about  200  feet 
from  the  shore,  in  the  N.  E.  ^  of  the  S.  E.  ^  of  S.  E.  ^  of 
Sec.  33,  T.  64  N.,  R.  8  W.  The  size  is  50x75  feet.  The 
surface  is  rocky,  with  a  few  small  shrubs  growing  in  crevices, 
and  is  literally  covered  with  rock  lichens,  and  a  few  Cladonias 
and  Stereocaulons  growing  along  crevices  and  beginning  to 
spread  in  one  or  two  places.  The  following  species  were  easily 
detected. 

1.  Cladonia  rangiferina  (L.)  HOFFM. 

2.  Cladonia  rangiferina  (L.)  HOFFM.  var.  sylvatica  L. 

3.  Cladonia  pyxidata  (L.)  FR. 

4.  Cladonia  uncialis  (L.)  FR. 

5.  Cladonia  furcata  (Huos.)  FR. 


224  MINNESOTA    BOTANICAL    STUDIES. 

6.  Steieocaulon  paschale  (L.)  FR. 

7.  Umbilicaria  muhlenbergii  (Acn.)  TUCK. 

8.  Umbilicaria  pustulata  (L.)  HOFFM. 

9.  Endocarpon  fluviatile  DC. 

10.  Parmelia  conspersa(EHRH.)  ACH. 
n.  Parmelia  saxatilis  (L.)  FR. 

12.  Parmelia  caperata  (L.)  ACH. 

13.  Physcia  sp. 

14.  Physcia  stellaris  (L.)  TUCK. 

15.  Physcia  csesia  (HOFFM.)  NYL. 

16.  Leptogium  lacerum  (Sw.)  FR. 

17.  Ephebe  pubescens  FR. 

18.  Ephebe  solida  BORN. 

19.  Pannaria  microphylla  (Sw.)  DELIS. 

20.  Urceolaria  scruposa  (L.)  NYL. 

21.  Placodium  vitellinum  (EHRH  )  NAEG.  and  HEPP. 

22.  Rinodina  oreina  (AcH.)  MASS. 

23.  Lecanora  rubina  (VILL.)  ACH. 

24.  Lecanora  cinerea  (L.)  SOMMERF. 

25.  Buellia  petraea  (FLOT.,  KOERB.)  TUCK. 

Comparing  the  lichens  easily  detected  on  islands  numbers  one 
and  three — those  which  give  character  to  the  flora — whatever  rare 
species  may  have  escaped  notice,  we  find  that,  of  a  total  of  23 
species  and  varieties  on  the  first  and  25  on  the  second,  19  are  com- 
mon to  both  islands,  separated  by  several  miles.  The  lists  as  a 
whole  show  a  large  number  of  foliaceous  and  fruticulose  species  ; 
and  we  evidently  have  not  the  primitive  post-pleistocene  lichen 
population  of  these  rocky  islands,  which  indeed  must  have  dis- 
appeared centuries  ago.  It  is  the  more  remarkable  that  practic- 
ally the  same  species  have  succeeded  in  replacing  a  former 
flora  on  the  two  islands.  I  regret  that  time  was  wanting  for  the 
study  of  more  of  these  islands,  and  especially  of  some  farther 
from  the  shore  line.  The  growth  of  larger  forms  of  vegetation 
is  probably  beginning  to  effect  a  decrease  in  lichen  species  on 
island  number  one  for  otherwise,  being  larger,  it  should  have 


Fink :   LICHENS  OF  THE  LAKE  SUPERIOR  REGION.  225 

furnished  more  species  than  number  three  rather  than  a  smaller 
number.  But  it  was  only  on  number  two  that  we  found  the  un- 
mistakable evidence  of  the  effects  of  the  arboreal  vegetation 
in  exterminating  the  lichens.  Here  too  the  species  existing  are 
all  but  two  the  same  as  those  found  on  one  or  both  of  the  other 
islands,  but  the  number  is  reduced  to  little  more  than  one-third 
as  many  as  occur  on  either  of  them. 

The  succession  of  species  is  as  apparent  upon  trees  as  upon 
rocks  and  is  constantly  in  evidence  in  this  largely  undisturbed 
region  where  trees  of  various  ages  grow  side  by  side.  Some 
of  the  crustaceous  lichens,  of  such  genera  as  Pyrenula,  Ar- 
thonia  and  Graphis^  were  usually  found  on  young  trees  with 
smooth  bark.  As  the  substratum  becomes  more  rugged  with 
the  increasing  age  of  the  tree,  these  are  gradually  replaced  by 
foliaceous  and  fruticulose  species  as  Ramalinas ,  Usneas,  Par- 
mclias,  etc.  Finally  as  the  trees  die  certain  species  of  Calicium 
Cladonia,Peltigera,  Parmelia,  etc.,  become  the  dominant  types. 
It  is  not  possible,  nor  is  it  necessary  here,  to  give  a  detailed  ac- 
count of  relation  between  each  epiphytic  lichen  and  its  host, 
but  a  few  of  the  most  apparent  relationships  are  in  order. 
Acer  spicatum  Lam.  supports  A rthonia  dispersa  (Schrad.)  Nyl. 
over  the  whole  area.  Populus  tremuloides  Michx.  and  P.  bal- 
sa ntif  era  L.  bear  Pyrenula  leucoplaca  (Wallr.)  Kbr.  commonly. 
Some  conifers,  as  Pinus  resinosus  Ait.,  P.  strobus  L.,  Thuja 
occidentalis  L.,  serve  for  substrata  for  those  species  of  the  genus 
Calicium  which  grow  on  living  trees.  The  most  luxuriant 
growths  of  Usnea  were  found  on  Picea  mariana  (Mill.)  and 
Abies  balsamea  (L.)  Mill.  Gr aphis  scripta  (L)  Ach.  var.  recta 
(Humb.)  Nyl.  was  almost  wholly  confined  to  Betula  lutea 
Michx.  and  this  same  tree  also  supports  Sagedia  oxyspora 
(Nyl.)  Tuck  and  two  or  three  Pyrenulas. 

A  close  analysis  of  the  distribution  of  species  within  the  area 
studied  reveals  much  of  interest  even  though  it  is  a  rather  a  re- 
stricted region.  Of  the  258  species  and  varieties  listed,  96  were 
found  only  in  one  place,  32  in  two,  31  in  three,  and  the  remain- 
ing 99  were  collected  along  lake  Superior  and  also  inland,  in 
four  or  more  localities  and  are  known  to  be  generally  distributed 
over  the  whole  of  the  two  counties.  Also  of  those  found  in 
in  two  or  three  localities,  34  species  were  collected  at  some 
point  along  lake  Superior  and  also  beyond  the  ridge  of  high 
land  formed  by  the  Mesabi  range  and  the  Misquah  hills  and 


226  MINNESOTA    BOTANICAL    STUDIES. 

are  doubtless  quite  generally  distributed  over  the  territory  sur- 
veyed. Of  these  found  in  two  or  three  places,  13  more  were 
found  along  the  lake  and  inland,  but  none  beyond  the  divide 
mentioned  above.  These  are  doubtless  generally  distributed 
between  this  highest  land  and  lake  Superior,  and  of  course  may 
occur  northwest  of  this  high  area  as  well. 

Of  course  the  133  or  more  species  most  generally  distributed 
over  the  area  largely  determined  the  character  of  its  flora  and 
are  interesting  in  studving  the  relation  of  the  flora  of  the  region 
to  that  of  others.  But  for  the  study  of  distribution  within  the 
area,  as  influenced  by  natural  conditions,  the  chief  interest  cen- 
ters around  the  96  species  found  in  one  place  only  and  those 
found  in  two  or  three  areas  and  yet  not  generally  distributed 
over  the  whole  region.  I  give  below  a  list  stating  the  whole 
number  collected  in  each  principal  collecting  ground  and  also 
the  number  found  at  each  one  and  not  elsewhere.  It  will  readily 
be  seen  that  the  last  datum  for  each  locality  simply  bears  a  close 
relative  proportion  to  the  first,  or  in  other  words,  that  no  one 
area  shows  a  large  relative  proportion  of  the  rare  species.  Of 
course  the  data  as  to  occurrence  of  these  rare  lichens  can  not  be 
relied  on  fully ;  but  about  two-thirds  of  them  are  species  of  size 
large  enough  to  be  easy  of  detection,  and  while  these  may  occur 
in  other  places,  they  are  surely  not  common  in  the  area.  The 
table  of  species  is  as  follows  : 

Grand  Portage  island,  total  collected,     88,  not  found  elsewhere,      15 

Grand  Portage,  "  "  59,  "  "  "  12 

South  Fowl  lake,  "  "  14,  "  "  "  i 

Rose  lake,  "•  "  20,  "  "  "  3 

Gunflint,  "  "  118,  "  "  "  12 

Misquah  hills,  "  "  115,  "  "  "  n 

Tofte,  "  "  85,  "  "  "  8 

Beaver  Bay,  "  "  82,  "  "  "  13 

Palisades,  "  "  33,  "  "  "  i 

Two  Harbors,  "  "  14,  k'  "  "  2 

Ely,  "  "  41,  «  "  "  3 

Snowbank  lake  area,  "  "  121,  "  "  "  15 

The  rather  high  per  cent,  of  forms  collected  at  Grand  Port- 
age only  is  due  to  the  fact  that  the  attempt  was  made  to  collect 
here  especially  species  not  found  on  the  island.  At  an  average 
about  one-eighth  of  the  species  collected  in  each  locality  were 
not  found  elsewere.  As  stated  above,  two-thirds  of  these  are 


Fink  :     LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  227 

conspicuous  forms.  The  remaining  one-third  are  some  of  the 
less  conspicuous  Biatoras,  Lccidcas,  Buellias^  Graphtses, 
Pyrenulas,  etc.,  which  are  not  easily  found. 

When  we  consider  the  limited  size  of  the  area  studied,  the 
restriction  of  certain  of  the  rarer  species  to  certain  parts  of  it 
rather  than  to  others  is  worthy  of  careful  study.  The  area  lies 
on  two  sides  of  a  divide  extending  approximately  east  and  west 
and  formed  of  the  Mesabi  range  and  the  Misquah  hills.  The 
alpine  or  sub-alpine  species  not  generally  distributed  over  the 
area  are  mostly  confined  to  the  portion  lying  between  the  divide 
and  lake  Superior  and  to  the  Misquah  hills  on  the  divide,  and 
also  those  found  along  the  lake  are  for  the  most  part  found 
toward  the  northeastern  portion  of  the  territory  traversed.  The 
rarer  temperate  region  species  on  the  other  hand  are  most 
numerous  to  the  north  and  west  of  the  divide,  or  toward  the 
south  of  the  portion  between  the  lake  and  the  divide.  All  of 
the  data  given  above  as  to  distribution  within  the  area  studied 
are  based  upon  carefully  prepared  lists  showing  the  distribution 
of  each  species.  They  can  not  all  be  reproduced,  but  parts  of 
them  must  be.  First  of  all,  the  facts  concerning  the  species 
found  only  in  one  place  can  only  be  properly  presented  for  con- 
sideration by  the  somewhat  laborious  table  below,  giving  the 
various  localities  and  species  for  each. 

GRAND  PORTAGE  ISLAND. 
Usnea  cavernosa  TUCK.,  N. 
Physcia  hispida  (SCHREB.)  FR.,  N. 
Solorina  saccata  (L.)  ACH.,  N. 

Lecanora  calcarea  (L.)  SOMMERF.  var.  contorta  FR.,  T. 
Cladonia  gracilis  (L.)  NYL.  var.  symphycarpia  TUCK.,  T. 
Cladonia  squamosa  HOFFM.  var.  phyllocoma  RABENH.,  T. 
Cladonia  deformis  (L.)  HOFFM.,  N. 
Baeomyces  byssoides  (L.)  SCHAER.,  N. 
Biatora  turgida  (FR.)  NYL.,  T. 
Lecidea  spirea  ACH.,  N. 
Endocarpon  miniatum  (L.)  SCHAER.,  T. 
Lecidea  crustulata  ACH.,  N. 
Lecidea  enteroleuca  FR.  var.  achrista  SOMMERF.,  T. 


228  MINNESOTA    BOTANICAL    STUDIES. 

Staurothele  drummondii  TUCK.,  T. 
Pyrenula  cinerella  (FLOT.)  TUCK.,  T. 

GRAND  PORTAGE. 

Parmelia  perforata  (jAcoJ  ACH.  var.  hypotropa  NYL.,  T. 
Physcia  adglutinata  (FLOERK.)  NYL.,  T. 
Umbilicaria  hyperborea  HOFFM.,  N. 
Nephroma  laevigatum  ACH.  var.  parile  NYL.,  N. 
Placodium  murorum  (HOFFM.)  DC.  var.  miniatumTucK.,  T. 
Lecanora  muralis  (SCHREB.)  SCHAER  var.  diffracta  FR.,  T. 
Biatora  leucophaea  FLOERK.  var.  griseoatra  KOERB.,  N. 
Biatora  lucida  (ACH.)  FR.,  N. 
Lecidea  lapicida  FR.  var.  oxydata  FR.,  N. 
Thelocarpon  prasinellum  NYL.,  T. 
Verrucaria  nigrescens  PERS.,  T. 

Pyrenula  cinerella  (FLOT.)  TUCK.  var.  quadriloculata  var. 
nov.  (?). 

SOUTH  FOWL  LAKE. 

Alectoria  saepincola  (EHRH.)  ACH.,  N. 

ROSE  LAKE. 

Heterothecium  sanguinarium  (L.)  FLOT.  var.  affine  TUCK.,  N. 
Opegrapha  varia  (PERS.)  FR.  var.  notha  ACH.,  T. 
Usnea  barbata   (L.)  FR.  var.  dasyopoga  FR.,  T. 

GUNFLINT. 

Physcia  caesia  (HOFFM.)  NYL.,  T. 
Pannaria  nigra  (Huos.)  NYL.,  T. 
Placodium  cinnabarrinum  (Acn.)  ANZ.,  T. 
Lecanora  pallida  (SCHREB.)  SCHAER.,  T. 
Lecanora  hageni  ACH.,  T. 

Cladonia  symphycarpa  FR.  var.  epiphylla  (Acn.).  NYL.,  T. 
Cladonia  fimbriata  (L.)  FR.,  T. 

Cladonia  gracilis  (L.)  NYL.  var.  cervicornis  FLOERK.,  T. 
Biatora  glauconigrans  TUCK.,  T. 


Fink :   LICHENS  OF  THE  LAKE  SUPERIOR  REGION.          229 

Lecidea  acclinis  FLOT.,  T. 

Buellia  petraea  (FLOT.,  KOERB.)  TucK.var.  grandis  FLOERK., 
N. 
Arthonia  patellulata  NYL.,  T. 

MlSQUAH    HILLS. 

Parmelia  centrifuga  (L.)  ACH.,  N. 

Collema  flaccidum  ACH.,  T. 

Placodium  murorum  (HOFFM.)  DC.,  T. 

Lecanora  subfusca  (L.)  ACH.  var.  hypnorum  SCHAER.,  T. 

Pertusaria  glomerata  (Acn.)  SCHAER.,  N. 

Biatora  oxyspora  (TUL.)  NYL.,  T. 

Biatora  schweinitzii  FR.,  T. 

Lecidea  lapicida  FR.,  N. 

Lecidea  albocaerulescens   WULF.)  SCHAER.,  N. 

Lecidea  platycarpa  ACH.,  N. 

Graphis  scripta  (L.)  ACH.  var.  limitata  ACH.,  T. 

TOFTE. 

Parmelia  perforata  QACO^)  ACH.,  T. 
Sticta  limbata  (SM.)  ACH.,  N. 
Leptogium  myochroum  (EHRH.)  SCHAER.,  T. 
Lecanora  elatina  ACH.,  T. 
Stereocaulon  coralloides  FR.,  N. 
Cladonia  caespiticia  (PERS.)  FL.,  T. 
Cladonia  digitata(L.)  HOFFM.,  N. 
Calicium  chrysocephalum  (TURN.)  ACH.  var.  filare  ACH.,  T. 

BEAVER  BAY. 

Ramalina  calicaris  (L.)  FR.  var.  fastigiata  FR.,  T. 
Ramalina  pollinarella  NYL.,  T. 

Placodium  cerinum(HED\v.)  NAEG.  and  HEPP.  var.  pyracea 
NYL.,T. 

Placodium  vitellinum  (EHRH.)  NAEG.  and  HEPP.,  T. 
Lecanora  calcarea  (L.)  SOMMERF.,  T. 


230  MINNESOTA    BOTANICAL    STUDIES. 

Cladonia  mitrula  TUCK.,  T. 
Cladonia  delicata  (EHRH.)  FL.,  T. 
Cladonia  decorticata  FLOERK.,  T. 
Biatora  coarctata  (SM.,  NYL.)  TUCK.,  T. 
Biatora  myriocarpoides  (NYL.)  TUCK.,  T. 
Biatora  naegelii  HEPP.,  T. 
Buellia  myriocarpa.  (DC.)  MUDD.,  T. 
Sagedia  oxyspora  (NYL.)  TUCK.,  T. 

PALISADES. 
Cetraria  islandica  (L.)  ACH.,  N. 

Two  HARBORS. 

Physcia  aquila  (AcH.)  NYL.,  T. 
Buellia  dialyta  (NYL.)  TUCK.,  T. 

ELY. 

Lecanora  tartarea  (L.)  ACH.,  T. 

Buellia  myriocarpa  (DC.)  MUDD.  var.  polyspora  WILLEY.,  T. 
Calicium  trichiale  (Acn.)  var.  stemoneum  NYL.  T. 

SNOWBANK  LAKE  AREA. 

Ramalina  calicaris  (L.)  FR.  var.  canaliculata  FR.,  T. 
Ramalina  pusilla  (PREV.)  TUCK.,  N. 

Parmeliatiliacea(HoFFM.)FLOERK.  var.  sublaevigataNYL.,T. 
Collema  pycnocarpum  NYL.,  T. 
Leptogium  lacerum  (Sw.)  FR.,  T. 

Leptogium  lacerum  (Sw.)  FR.  var.  pulvinatum   MOUG.  and 
NESTL.,  T. 

Rinodina  sophodes  (Acn.)  NYL.  var.  confragosa  NYL.,  T. 

Gyalecta  fagicola  (HEPP.)  TUCK.,  T. 

Biatora  sphaeroides  (DICKS.)  TUCK.,  T. 

Biatora  fuscorubella  (HOFFM.)  TUCK.,  T. 

Biatora  muscorum  (Sw.)  TUCK.,  T. 

Lecidea  cyrtidia  TUCK.,  T. 

Buellia  parmeliarum  (SOMMERF.)  TUCK.,  T. 


Fink  :     LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  231 


Coniocybe  pallida  (PERS.)  FR.,  T. 
Verrucaria  epigsea  (PERS.)  Acn.,  T. 


In  the  above  table  all  of  the  arctic  and  subarctic  species  are 
marked  (N.)  and  the  species  common  in  temperate  regions  at 
low  elevations  (T.).  Beginning  then  with  the  areas  between 
the  divide  formed  by  the  Mesabi  range  and  the  Misquah  hills 
and  lake  Superior  and  toward  the  north  of  this  region,  of  a 
total  of  15  species  found  only  on  Grand  Portage  island,  seven 
or  one  less  than  half  are  characteristic  of  northern  regions. 
Of  12  confined  to  Grand  Portage,  five  or  about  42  per  cent,  are 
northern  species.  A  single  one  collected  only  at  South  Fowl 
lake  and  one  of  three  at  Rose  lake  are  also  northern.  Of  the 
12  found  only  at  Gunflint  and  the  15  found  only  in  the  Snow- 
bank lake  area,  only  one  strictly  northern  species  is  restricted 
to  each  place,  while  for  Ely  of  the  three  restricted  species  not 
one  is  northern.  Consideration  of  the  figures  will  show  that  for 
the  five  localities  along  the  international  boundary  there  is  a  de- 
crease in  proportion  of  rare  arctic  or  alpine  species  in  passing 
westward  along  the  boundary.  Again  passing  southward  from 
Grand  Portage  we  found  three  northern  of  a  total  eight  species 
confined  to  Tofte,  and  the  one  species  taken  only  at  the  Pali- 
sades is  northern.  The  thirteen  species  found  only  at  Beaver 
Bay  and  the  two  collected  only  at  Two  Harbors  are  all  distinctly 
temperate  region  plants.  Thus  it  appears  that  the  northern 
species  give  way  to  those  more  characteristic  of  temperate  re- 
gions in  passing  southward  even  along  the  shores  of  lake  Su- 
perior where  the  cold  lake  winds  have  greatest  influence  upon 
the  flora.  An  elevation  of  more  than  1,000  feet  was  reached 
at  Beaver  Bay  without  finding  northern  species  while  at  Grand 
Portage  about  150  miles  northeast  they  descend  to  the  lake  level. 
At  Tofte  a  short  distance  northeast  of  Beaver  Bay  we  reached 
an  elevation  of  1,529  feet  on  Carlton  peak  in  the  Sawteeth 
mountains  and  found  three  northern  species.  Only  one  of  the 
three  species  was  taken  at  the  summit  of  Carlton  peak,  but  this 
is  because  the  top  of  the  peak  is  burned  over.  The  other  two 
species  were  collected  at  a  considerable  distance  above  the  base 
of  the  peak  and  are  doubtless  to  be  found  on  unburned  portions 
of  the  Sawteeth  mountains  near  by. 

In  the  discussion  of  the  table  thus  far  the  Misquah  hills  area 
has  not  been  considered.  As  stated  elsewhere  this  area  is  the 


232  MINNESOTA    BOTANICAL    STUDIES. 

highest  portion  of  the  State,  reaching  2,230  feet,  and  the  region 
gave  five  arctic  or  subarctic  species  of  a  total  of  eleven  collected 
only  here.  The  influence  of  elevation  becomes  apparent  when 
we  compare  the  Misquah  hills  area  with  the  Gunflint  region  as 
to  number  of  northern  species.  The  latter  locality  lies  20  miles 
north  of  the  former  and  about  400  feet  lower,  only  a  small  por- 
tion of  it  near  the  Paulson  mines  reaching  2,000  feet.  The 
Misquah  hill  area  which  is  a  more  extended  region  of  higher 
elevation  furnished  the  goodly  proportion  of  northern  forms 
noted  above  while  the  Gunflint  area  gave  only  one  such  species 
in  a  total  of  twelve  found  only  in  the  area.  . 

Of  all  the  species  found  only  in  one  place  25,  or  more  than 
one-fourth,  are  arctic  or  subartic  and  71,  or  approximately 
three-fourths  are  plants  characteristic  of  temperate  regions. 
Without  entering  into  the  yet  more  complicated  analysis  which 
a  consideration  of  these  rare  temperate  region  plants  would 
involve,  a  mere  inspection  of  the  table  will  show  in  a  general 
way  that  their  distribution  is  just  the  reverse  of  that  of  the  north- 
ern species,  or  that  they  are  especially  characteristic  of  that 
small  portion  of  the  region  studied  which  lies  to  the  north  of  the 
divide  and  of  the  southern  portion  of  the  region  lying  between 
the  divide  and  lake  Superior.  Of  course  it  could  also  be 
shown  that  they  are  more  especially  characteristic  of  the  lower 
elevations. 

Of  the  63  species  collected  only  in  two  or  three  places,  only 
a  half  dozen  are  arctic  or  subarctic  species,  and  it  would  have 
been  useless  to  give  the  whole  63  in  tabular  form  as  no  safe 
data  could  be  obtained  from  so  small  a  proportion  of  northern 
species.  However,  the  6  northern  species  are  as  follows  : 

Ramalina  pusilla  (PREV.)  TUCK.  var.  geniculata  TUCK. 

Parmelia  encausta  (SM.)  NYL. 

Sticta  scorbiculata  (Scop.)  ACH. 

Lecanora  frustulosa  (DICKS.)  MASS. 

Lecanora  sordida  (PERS.)  TH.  FR. 

Buellia  geographica  (PERS.)  TUCK. 

In  order  that  we  may  have  all  of  the  northern  species  before 
us  for  a  final  consideration,  I  shall  give  a  list  of  those  generally 
distributed  as  follows  : 

Lecidea  lactea  FL. 


Fink :   LICHENS  OF  THE  LAKE  SUPERIOR  REGION.          233 

Buellia  petraea  (PLOT.,  KOERB.)  TUCK. 

Buellia  petraea  (PLOT.,  KOERB.)  TUCK.  var.  montagnaei  TUCK. 

Umbilicaria  vellea  (L.)  NYL. 

Nephroma  tomentosum  (HOFFM.)  NECK. 

Pannaria   epidiota  TH.  PR. 

Stereocaulon  paschale  (L.)  PR. 

Cladonia  amaurocrasa  (PL.)  SCHAER. 

Baeomyces  aeruginosus  (Scop.)  DC. 

Heterothecium  sanguinarium  (L.)  PLOT. 

Taking  into  account  the  above  table,  we  find  that  of  a  total 
of  99  species  and  varieties  generally  distributed  over  the  area 
studied  only  10,  or  one-ninth,  are  arctic  or  subarctic,  and  con- 
sidering both  of  the  last  two  tables  we  see  that  of  162  species 
and  varieties  more  or  less  widely  distributed  16,  or  nearly  one- 
ninth,  are  arctic  or  subarctic.  It  has  been  stated  that  about 
one-fourth  of  the  species  found  only  in  one  place  are  such  north- 
ern forms.  Thus  we  find  that  the  more  general  the  distribution 
of  a  series  of  plants  in  the  area  the  smaller  the  per  cent,  of 
northern  species,  and  conversely  the  larger  the  per  cent,  of 
temperate  region  species.  In  other  words  the  prevailing  species 
are  those  characteristic  of  temperate  regions,  and  as  a  whole  the 
rarer  ones  are  the  more  northern  floral  elements.  Since  the  in- 
troduction of  new  species  is  commonly  a  more  rapid  process 
than  the  complete  extermination  of  others  in  a  given  region,  the 
existing  conditions  above  stated  seem  to  prove,  as  one  would 
naturally  suppose,  that  the  present  lichen  flora  of  the  region  is  in 
general  of  temperate  region  elements  and  that  the  more  northern 
elements  of  the  flora  are  the  persisting  for  most  part  in  a  few  favor- 
able spots.  This  supposition  also  explains  the  existence  of  the 
northern  species  in  isolated  regions  further  south  as  I  have  done 
for  Taylors  Falls.  Professor  Conway  MacMillan  has  con- 
sidered the  spermaphytic  flora  of  this  region  as  a  south-bound 
one,*  or  at  least  that  of  the  portion  between  the  divide  and  lake 
Superior.  My  observations  here  and  at  Taylors  Falls  do  not 
indicate  that  this  is  generally  true  of  the  lichens.  However, 
because  of  somewhat  milder  temperature,  lower  elevation  and 
perhaps  more  early  retreat  of  the  ice  sheet  in  the  western  half 

*  MacMillan  C.  Observations  on  the  distribution  of  plants  along  the  shore 
at  Lake  of  the  Woods.  Minn.  Bot.  Stud,  i  :  954.  1897. 


234  MINNESOTA    BOTANICAL    STUDIES. 

of  the  State,  the  temperate  region  lichens  have  no  doubt  driven 
or  followed  the  northern  species  farther  north  there  and  doubt- 
less very  few  of  the  latter  elements  now  exist  in  western  Min- 
nesota south  of  Lake  of  the  Woods,  or  indeed  anywhere  in  the 
western  half  of  the  state.  Since  gaining  possession  of  the  north- 
western part  of  the  state  these  temperate  region  lichens  have 
doubtless  been  moving  southward  over  the  Mesabi  range  to 
meet  similar  floral  elements  of  a  generally  northward  bound 
lichen  flora.  The  western  half  of  the  state  remains  to  be  ex- 
plored for  lichens,  but  the  above  statement  as  to  the  character 
of  the  flora,  based  on  observations  recorded  in  this  paper  for 
localities  north  of  the  Mesabi  range  and  some  knowledge  of  the 
general  character  of  the  region,  I  regard  as  sufficiently 
secure. 

Like  the  Taylors  Falls  region  this  one  of  course  at  one  time 
contained  only  arctic  species,  and  the  present  more  numerous 
species  characteristic  of  temperate  regions  have  gained  the 
ascendency  in  quite  recent  time.  However,  the  problems  in- 
volved in  the  struggle  betwen  the  contending  floral  elements 
do  not  force  themselves  upon  the  observer  so  strongly  in  this 
larger  area  and  must  be  dismissed  with  a  much  briefer  state- 
ment. As  in  the  Taylors  Falls  region  the  persisting  northern 
species  are  largely  lithophytic.  This  is  shown  in  the  following 
exhibit  of  substrata  for  the  42  species  : 

Arctic  or  subarctic  lichens  confined  to  rocks 25 

"  "  "  "  "  "  trees S 

"  "  "  "  "  "  earth 4 

"  "  "  "  "  "  dead  wood 2 

"  "  "  "  "  "  dead  wood  and  trees  i 

"  "  "  u  "  "   earth  and  rocks I 

"  "  "  "  "  "  trees     "        "                  i 

The  greater  persistence  of  the  lithophytic  species  is  doubtless 
due  in  part  at  least  to  the  greater  stability  of  the  rock  surfaces 
and  also  probably  in  part  to  the  fact  that  the  arctic  and  sub- 
arctic species  became  more  thoroughly  established  on  the  rocks, 
which  were  present  for  them  to  attack  immediately  at  the  close 
of  the  Pleistocene  before  the  advent  of  large  trees  and  temper- 
ate-region lichens. 

Thus  far  I  have  given  a  detailed  account  of  habitat  for  the 
northern  species  only.  Of  the  whole  258  species  and  varieties 


Fink  :     LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  235 

listed  in  this  paper,  so  far  as  observation  showed,  83  are  epi- 
phytic, 80  lithophytic  and  29  may  occur  on  either  trees  or  rocks. 
Another  29  are  epigean,  22  were  found  only  on  dead  wood,  six 
are  lithophytic  or  epigean,  three  are  found  on  earth  and  dead 
wood,  three  are  parasitic  on  other  lichens  and  two  were  found 
on  living  and  dead  wood.  The  above  analysis  of  substrata  is 
somewhat  different  from  that  used  in  the  second  paper  of  this 
series  for  comparison  of  substrata  at  Minneapolis  and  at  Fayette, 
Iowa ;  but  when  reduced  to  that  form  shows  that  the  percentage 
of  lichens  growing  on  rocks  is  somewhat  higher  than  that  for 
the  two  areas  farther  south,  while  the  percentage  of  those  grow- 
ing on  wood  is  considerably  lower.  The  larger  proportion  of 
rock  lichens  in  the  Superior  region  is  due  to  at  least  three  things. 
First,  the  more  extensive  exposure  of  rock  surfaces,  though  this 
is  offset  in  part  at  least  by  the  existence  of  three  distinct  kinds 
of  rock  in  the  Minneapolis  region,  viz.  :  the  igneous  or  meta- 
morphic  boulders,  the  limestone  outcrops  and  the  Saint  Peter 
sandstone.  Second,  the  rock  surfaces  become  warmer,  each 
day  in  the  warm  portion  of  the  year,  than  the  trees,  because  of 
rapid  absorption  of  heat ;  and  this  doubtless  favors  lichen  de- 
velopment on  rocks  in  this  northern  region.  Third,  general 
moisture  of  much  of  the  surface  due  to  the  fact  that  the  rocks 
are  comparatively  impervious  to  water,  so  that  much  of  it  col- 
lects in  lakes  and  swamps,  favors  good  development  of  rock 
lichens  as  compared  with  the  region  about  Minneapolis.  At 
Minneapolis  unshaded  rocks  bear  very  few  lichens,  but  in  the 
Superior  region  rocks  are  well  populated  with  them  at  all  eleva- 
tions and  in  all  sorts  of  environment  at  or  above  the  water  line, 
except  where  killed  by  fire. 

The  total  number  of  genera  for  the  region  is  39,  while  the 
number  for  Minneapolis  and  Taylors  Falls,  so  far  as  is  known, 
is  34.  The  whole  number  for  Illinois,  as  given  by  Wolf  and 
Hall*  is  40  and  for  Iowa  as  recorded  by  the  writer  in  two  papersf 
is  38.  This  total  number  of  genera  for  the  Superior  region  is 
seen  by  the  comparisons  with  both  larger  and  smaller  areas 
further  south  to  be  rather  large  for  a  somewhat  limited  northern 


*  Wolf,  John  and  Hall,  Elihu.  A  List  of  the  Mosses,  Liverworts  and  Lich- 
ens of  Illinois.  Bull.  Ills.  State  Lab.  Nat.  Hist.  2  :  18-34.  June,  1878. 

fFink  B.  Lichens  of  Iowa.  Bull.  Lab.  of  Nat.  Hist.,  State  Univ.  of  Iowa  3: 
70-88.  Mr.  1895,  and  Notes  concerning  Iowa  Lichens,  Prgc.  Iowa  Acad.  Sci. 
5  :  174-187.  1897. 


236  MINNESOTA    BOTANICAL    STUDIES. 

area,  but  a  locality  where  northern  and  more  temperate  floral  ele- 
ments meet  seems  to  compensate  fully  at  least  for  difference  in 
latitude. 

The  genera  giving  most  of  the  species  new  to  the  state  are 
the  following,  given  in  tabular  form  with  the  total  number  of 
species  collected  in  each  genus  and  the  number  new  to  the 

state. 

Ramalina,    collected  6,  new  to  the  State  4. 

Cetaria,  "  7,  "  "  "  5. 

Sticta,  "  5,  "  "  "  3. 

Nephroma,  "  4,  "  "  "  3- 

Solorina,  "  i,  "  "  "  i. 

Lecanora,  "  25,  "  "  "  15. 

Gyalecta,  "  i,  "  "  "  i. 

Bceomyces,  "  2,  "  "  "  2. 

Biatora,  "  23,  "  "  "  16. 

Heterothecium,  "  2,  "  "  "  2. 

Lecidca,  "  12,  "  "  "  9. 

Calicium,  "  9,  "  "  "  9. 

Coniocybe,  "  i,  "  "  "  i. 

Sagedia,  "  i,  "  "  "  i. 

Of  these  genera  Solorina,  Bceomyces,  Heterothecium,  Calic- 
tumy  Coniocybe,  and  Sagedia  are  new  to  the  State.  An  inspec- 
tion of  the  above  list  shows  that  the  genera  are  for  the  most 
part  those  furnishing  large  numbers  of  arctic  and  subarctic 
species,  or  species  hitherto  supposed  to  be  confined  to  New 
England.  The  genera  Stereocaulon  and  Umbilicaria  are 
equally  characteristic  of  northern  and 'eastern  areas,  but  two- 
thirds  of  the  species  of  these  genera  here  recorded  were  listed 
for  Minnesota  in  the  first  paper  of  this  series. 

By  comparing  the  present  list  of  species  and  varieties  with 
those  recorded  for  Minneapolis  and  Taylors  Falls  we  find  that 
that  there  are  152  lichens  growing  in  the  territory  considered  in 
this  paper  and  not  found  in  either  of  the  two  areas  named  above, 
while  there  are  33  found  in  them  and  not  in  the  northeastern 
Minnesota  area  under  consideration.  This  leaves  only  73 
lichens  known  to  be  common  to  central  and  northeastern  Min- 
nesota. In  the  comparison  between  Minneapolis  and  certain 
localities  in  northeastern  Iowa  it  was  shown  that  no  species  have 
been  found  at  the  former  place  and  not  in  the  latter  region, 
though  Minneapolis  is  150  miles  north  of  Fayette,  the  principal 
Iowa  area  considered.  In  passing  about  200  miles  north  from 


Fink:     LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  237 

Minneapolis,  on  the  other  hand,  we  find  a  lichen  flora,  about 
three-fifths  of  whose  species  and  varieties  are  not  found  at  Min- 
neapolis and  about  half  of  which  are  new  to  the  state.  The 
region  of  rapid  transition  in  lichen  species  lies  between  the 
Minneapolis  and  Superior  areas  and  has  only  been  touched  in 
the  study  of  its  rock  lichens  at  Taylors  Falls.  As  'stated  in  the 
third  paper  of  this  series,  this  region  is  one  of  special  interest 
for  tracing  the  distribution  of  species.  The  cause  of  the  great 
difference  in  lichen  flora  between  Minneapolis  and  the  Superior 
region  is  scarcely  due  in  any  great  measure  to  difference  in 
latitude  since  an  almost  equal  difference  in  latitude  to  the  south 
of  Minneapolis  caused  no  appreciable  difference  in  the  flora. 
Also,  I  have  shown  in  this  paper  that  in  three  localities  lying 
in  the  northern  part  of  the  region  studied  in  the  paper,  viz.  : 
Snowbank  lake,  Ely  and  Gunflint,  very  few  typically  northern 
species  are  found.  These  regions  at  the  north  of  the  area  are 
more  closely  related  to  the  Minneapolis  region  as  to  lichen  flora 
than  others  50  or  75  miles  further  south.  The  difference  in 
lichen  flora  between  central  and  northeastern  Minnesota  seems 
then  to  be  due  chiefly  to  three  factors.  The  first  is  difference 
in  substrata.  The  limestones  of  the  Minneapolis  regions,  as 
well  as  the  sandstone,  are  almost  entirely  wanting  in  northeast- 
ern Minnesota,  being  replaced  by  an  abundance  of  igneous  or 
metamorphic  rocks.  The  conifers,  which  abound  in  the  north- 
ern part  of  the  State,  and  which  serve  for  substrata  for  quite  a 
number  of  species  not  found  southward,  form  the  other  chief 
difference  as  to  substrata.  Location  in  the  valley  of  lake 
Superior,  where  the  region  is  shut  off  from  warmer  regions  west 
and  north  as  well  as  south,  is  another  factor  that  has  caused 
much  of  the  difference  in  flora.  It  has  been  shown  that  the 
number  of  arctic  and  subarctic  species  for  a  given  elevation 
decreases  in  passing  southwestward  along  the  lake.  This  I 
suppose  to  be  due  not  so  much  to  difference  in  latitude  as  to  the 
fact  that  in  the  northeastern  part  of  the  region  studied  along  the 
lake  the  winds  coming  from  the  broader  expanse  of  water,  on 
this  largest  American  fresh  water  area,  are  rendered  cooler  than 
farther  down  where  the  lake  is  not  so  wide.  That  the  cold 
winds  are  a  factor  is  demonstrated  by  the  occurrence  of  a  large 
proportion  of  northern  species  at  the  Grand  Portage  area,  and 
especially  on  the  island  which  rises  little  more  than  100  feet 
above  the  lake,  while  such  inland  areas  as  Gunflint  and  Snow- 


238  MINNESOTA    BOTANICAL    STUDIES. 

bank  lake,  which  are  somewhat  farther  north,  are  almost 
entirely  devoid  of  such  species.  A  third  factor  is  increase  in 
elevation.  The  influence  of  elevation  has  been  discussed  in 
considering  the  Misquah  hills  and  the  Sawteeth  mountains. 
Concerning  cold  lake  winds  and  elevation,  it  is  significant  that 
of  the  25  arctic  and  subarctic  species  found  only  in  one  place, 
16  are  found  at  stations  along  the  lake,  and  that  of  the  remain- 
ing 9,  5  are  found  in  the  Misquah  hills,  the  region  of  highest 
elevation.  Thus  all  but  4  of  these  25  species  were  collected 
where  one  or  both  of  these  factors  have  most  influence. 

It  is  not  possible  to  state  just  which  ones  of  the  many  species 
found  either  in  central  or  in  northeastern  Minnesota  and  not  in 
the  other  area  would  be  of  most  interest  in  studying  the  territory 
lying  between  the  two  regions.  Of  course,  the  foliaceous  and 
fruticulose  species  are  most  easily  found,  and  some  of  these  are 
most  likely  to  be  collected.  I  may  add  that  interest  would 
centre  chiefly  about  species  which  are  common  and  give  char- 
acter to  the  flora  in  one  of  the  two  areas  and  are  not  found  in 
the  other.  Not  attempting  to  select  from  some  150  species  col- 
lected in  the  Superior  area  and  not  farther  south  in  the  state,  I 
will  name,  from  27  or  28  species  found  to  the  south  and  not  to 
the  north  of  the  unexplored  area,  Theloschistes  concolor  (Dicks.) 
Tuck,  and  Physcia  granulifera  (Ach.)  Tuck,  as  two  species 
that  any  botanist  can  soon  learn  to  distinguish  in  the  field, 
which  are  common  in  the  south  half  of  Minnesota  and  not 
known  farther  north  in  the  state,  and  whose  distribution  be- 
tween Minneapolis  and  Two  Harbors  would  be  of  special  inter- 
est. Nearly  all  of  the  remainder  of  the  27  or  28  species  are 
either  infrequent  or  rare  about  Minneapolis,  are  confined  to  sub- 
strata not  existing  in  northeastern  Minnesota,  or  are  so  incon- 
spicuous as  to  render  their  study  in  the  field  difficult. 

As  a  fitting  close  to  these  observations  on  the  distribution  of 
lichens  in  different  regions  of  the  state  I  may  give  some  notes 
concerning  certain  species  for  most  part  characteristic  of  more 
southern  portions  and  found  also  in  northeastern  Minnesota. 
Parmelia  borreri  Turn,  was  not  found  fruited  along  the  lake 
north  of  Beaver  Bay.  Peltigera  aphlhosa  (L.)  Hoffm.  seeks 
high  ground  in  the  southern  stations  from  which  it  is  recorded 
and  in  the  northwestern  as  well  and  is  one  of  the  species  whose 
farther  southern  and  western  extent  in  the  state  would  be  espe- 
cially worthy  of  study.  Parmelia  tiliacea  (Hoffm.)  Tuck,  be- 


Fink  :     LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  239 

comes  somewhat  common  at  the  southwest  and  more  so  at  the 
northwest  portion  of  the  territory  studied.  It  is  one  of  the 
southern  intrusions  which  extend  farther  north  at  some  distance 
from  lake  Superior.  Trees  common  farther  south  in  Min- 
nesota were  noted  north  of  the  Mesabi  range  as  Qiiercus  macro- 
carpa  Michx.  Cratcegus  sp.  and  Fraxinus  sp.  On  Fraximis 
was  found  Coniocybe  -pallida  (Pers.)  Fr.  which  is  common  in 
northeastern  Iowa,  and  on  the  same  host  Pyrenula  leucophaca 
(Wallr.)  Kbr.  also  becomes  common  for  the  first  time  in  this 
northern  area.  Other  lichens  in  the  list  of  those  found  only 
north  of  the  Mesabi  range  or  the  Misquah  hills  could  be  se- 
lected for  special  treatment,  and  on  the  whole  this  portion  of 
the  Superior  region  shows  a  closer  floral  connection  with  central 
Minnesota  than  does  the  most  southern  point  reached,  viz.,  Two 
Harbors. 

The  list  of  258  species  and  varieties  is  a  large  one  for  a 
rather  limited  area  to  yield,  especially  when  it  is  stated  that  only 
about  300  lichens  have  been  listed  for  Minnesota,  including 
the  present  list.  Yet  the  undisturbed  portions  of  the  region 
are  more  remarkable  for  richness  in  individuals  than  for  large 
numbers  of  species.  The  Grand  Portage  area  gave  132  species 
and  varieties  and  the  Snowbank  lake  area  121.  These  two 
areas  are  the  ones  best  studied  and  are  perhaps  as  thoroughly 
explored  as  the  Minneapolis  area,  which  gave  113  species  and 
varieties.  Both  of  the  former  two  are  like  the  latter  small  areas 
and  the  comparison  seems  to  indicate  that  the  lake  Superior 
region  is,  area  for  area,  only  slightly  richer  in  species  than  the 
Minneapolis  region.  Professor  L.  H.  Bailey*  found  that  the 
species  of  Spermaphytes  and  higher  Archegoniates  of  the 
region  are  only  about  one-tenth  as  numerous  at  lake  Vermilion 
as  in  similar  areas  six  degrees  farther  south,  and  we  should  of 
course  expect  some  decrease  in  lichen  species  rather  than  an 
increase  in  passing  northward  in  the  state.  The  reverse  con- 
dition existing  is  largely  due  to  diverse  conditions  within  the 
region  as  to  elevation  and  temperature  and  as  to  surface  moisture, 
all  of  which  factors  have  been  duly  considered.  The  Snow- 
bank lake  area  studied  is  a  larger  one  than  the  Grand  Portage. 
About  equal  time  was  spent  at  the  two  places,  and  we  did  much 
more  of  the  time-consuming  quantity-collecting  at  Grand  Port- 

*  Bailey,  L>  H.,  in  Arthur,  J.  C.     Report  on  botanical  work  in  Minnesota  in 
1886.     Geol.  and  Nat.  Hist.  Surv.  of  Minn.,  Bull.  3:8.     i  O.,  1887. 


240  MINNESOTA    BOTANICAL    STUDIES. 

age.  Yet  fewer  forms  of  lichens  were  collected  about  Snow- 
bank lake.  This  seems  to  indicate  that  the  lake  shore  is  richer 
in  lichen  species  than  in  interior  areas  of  the  territory  studied. 

It  is  well  known  that  a  large  portion  of  the  species  of  lichens 
of  the  interior  of  North  America  are  those  found  also  in  regions 
bordering  upon  the  Atlantic  ocean  along  our  eastern  border. 
This  was  brought  out  by  the  writer  in  a  previous  paper,*  but 
all  the  species  recorded  in  that  paper  were  temperate  region 
lichens.  It  has  remained  for  the  present  paper  to  record  a  large 
number  of  more  northern  lichens  previously  for  the  most  part 
known  only  in  arctic  or  subarctic  regions,  or  descending  from 
mountains  farther  south  only  along  our  Atlantic  border. 

Of  the  258  species  and  varieties  listed  below  46  are  new  to 
the  North  American  interior  or  to  the  interior  of  the  United 
States,  and  of  these  six  are  new  west  and  north  of  New  Eng- 
land. In  treating  of  distribution  the  expression  "  the  interior  of 
North  America  "  means  the  area  lying  between  the  Appalachian 
system  of  mountains  on  the  east  and  the  Rocky  Mountains  on 
the  west.  A  number  of  species  noted  as  new  to  the  interior 
have  been  reported  from  New  York  or  Canada,  and  doubtless  a 
few  of  them  were  previously  found  a  short  distance  west  of  the 
Appalachian  system  of  mountains. 

LIST    OF    SPECIES    AND    VARIETIES. 

1.  Ramalina  calicaris  (L.)  FR.  var.  fastigiata  FR. 

On  trees,  rare.     Beaver  Bay,  July  13,  1897,  no.  677. 

2.  Ramalina  calicaris  (L.)  FR.  var,  canaliculata  FR. 

On  trees,  rare.     Snowbank  lake  area,  July  23,  1897,  no.  895. 
Not  previously  reported  from  Minnesota. 

3.  Ramalina  calicaris  (L.)  FR.  var.  farinacea  SCHAER. 

On  rocks,  common  or  frequent  over  whole  area  and  rarely 
found  on  trees  also.  Grand  Portage  island,  June  23,  1897,  no. 
106.  Gunflint,  June  30,  1897,  no.  257.  Misquah  hills,  July 
3,  1897,  no.  419,  and  July  7,  1897,  no.  539.  Tofte,  July  10, 
1897,  no.  624.  Palisades,  July  15,  1897,  no.  763.  Snowbank 
lake  area,  July  24,  1897,  no.  934. 

All  except  no.  763  were  lighter  colored  than  other  forms  of 
the  species.  No.  106  occasionally  and  no.  763  quite  commonly 

*Fink,  B.  Lichens  of  Iowa.  Bull.  Lab.  of  Nat.  Hist.,  State  Univ.  of 
Iowa.  3  :  70-78.  Mr.,  1895. 


Fink  :     LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  241 

are  irregularly  branched  and  with  dilated  terminal  soredia  like 
/?.  pollinaria  (Acn.)  TUCK. 

4.  Ramalina  pusilla  (PREV.)  TUCK. 

On  trees,  frequent.  Grand  Portage  island,  June  19,  1897, 
no.  34.  Snowbank  lake  area,  July  20,  1897,  no.  844. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

5.  Ramalina  pusilla  (PREV.)  TUCK.  var.  geniculata  TUCK. 

On  trees,  infrequent  or  rare.  Gunflint,  July  2,  1897,  no.  375. 
Beaver  Bay,  July  13,  1897,  no.  675. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  the  United  States. 

6.  Ramalina  pollinarella  NYL. 

On  rocks,  rare.     Beaver  Bay,  July  13,  1897,  no.  681. 
Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior  of  North  America. 

7.  Cetraria  aurescens  TUCK. 

On  trees,  rare.  Tofte,  July  10,  1897,  no.  636.  Snowbank 
lake  area,  July  21,  1897,  no.  869. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

8.  Cetraria  islandica  (L.)  ACH. 

On  earth  above  the  Palisades,  rare,  July  15,  1897,  no.  765. 
Not  previously  reported  from  Minnesota. 

9.  Cetraria  ciliaris  (Acn.)  TUCK. 

On  trees,  abundant  on  Grand  Portage  island,  elsewhere  only 
rare  or  frequent.  Grand  Portage  island,  June  18,  1897,  no.  10. 
Gunflint,  July  2,  1897,  nos.  387  and  396  at  Misquah  hills,  July 
3,  1897,  no.  427.  Snowbank  lake  area,  July  22,  1897,  no. 
883,  and  July  26,  1897,  no.  948.  Ely,  July  28,  1897,  no.  1022. 

10.  Cetraria  lacunosa  ACH. 

.  On  trees,  common.  Top  of  bluff  at  south  end  of  South  Fowl 
lake,  June  26,  1897,  no  206.  Gunflint,  July  2,  1897,  no.  401. 
Misquah  hills,  July  3,  1897,  no.  442.  Tofte  (Carlton  peak), 
July  10,  1897,  no.  556.  Beaver  Bay,  July  14,  1897,  no.  733. 
Snowbank  lake  area,  July  21,  1897,  no.  946. 

Not  noted  at  Grand  Portage  where  the  last  above  was  abun- 
dant, but  seeming  to  replace  it  in  part  elsewhere,  being  com- 
mon in  the  localities  noted  above. 


242  MINNESOTA    BOTANICAL    STUDIES. 

11.  Cetraria  juniperina  (L.)  ACH.  var.  pinastri  ACH. 

On  trees  and  old  logs.  Common  at  Grand  Portage  and  Mis- 
quah  hills,  elsewhere  infrequent  or  rare,  sterile.  Grand 
Portage  island,  June  18,  1897,  no.  15.  Gunflint,  June  30, 
1897,  no.  261.  Misquah  hills,  July  3,  1897,  no.  411.  Beaver 
Bay,  July  13,  1897,  no.  679.  Snowbank  lake  area,  July  21, 
1897,  no.  865.  Ely,  July  28,  1897,  no.  1016.  « 

Not  previously  reported  from  Minnesota  and  new  to  the  inte- 
rior of  North  America. 

12.  Cetraria  saepincola  (EHRH.)  ACH. 

On  trees,  rare.     South  Fowl  lake,  June  27,  1897,  no.  201. 
Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

13.  Evernia  furfuracea  (L.)  MANN. 

On  trees,  rare,  sterile.  Gunflint,  July  2,  1897,  no.  397. 
Misquah  hills,  July  3,  1897,  no.  434.  Tofte  (Carlton  peak), 
July  10,  1897,  no.  573. 

Not  previously  reported  from  Minnesota. 

14.  Evernia  prunastri  (L.)  ACH. 

On  trees,  common.  Only  seen  fertile  once.  Grand  Portage 
island,  June  22,  1897,  no.  81.  Gunflint,  June  30,  1897,  no. 
267.  Misquah  hills,  July  3,  1897,  no.  428.  Tofte  (Carlton 
peak),  July  10,  1897,  no.  554.  Tofte,  July  10,  1897,  no.  635. 
Beaver  Bay,  July  14,  1897,  no.  717.  Two  Harbors,  July  17, 
1897,  no.  789.  Snowbank  lake  area,  July  20,  1897,  no.  841. 
Ely,  July  28,  1897,  no.  1002. 

15.  Usnea  barbata  (L.)  FR.  var.  florida  FR. 

On  trees,  common  or  abundant  but  sterile.  Grand  Portage 
island,  June  17,  1897,  no.  8.  Misquah  hills,  July  3,  1897,  no. 
420.  Tofte  (Carlton  peak),  July  10,  1897,  nos.  557  and  607. 
Beaver  Bay,  July  14,  1897,  no.  722.  Snowbank  lake  area, 
July  14,  1897,  No.  814. 

The  last  has  the  minute  and  numerous  fibrils  of  var.  hirta  Fr., 
but  it  is  not  sorediate ;  while  the  forms  given  below  under  that 
variety  are  sorediate,  but  the  fibrils  are  seldom  minute. 

16.  Usnea  barbata  (L.)  FR.  var.  hirta  FR. 

On  trees,  common  or  abundant  but  sterile.  Grand  Portage 
island,  June  16,  1897,  no.  9.  Gunflint,  July  i,  1897,  no.  356. 
Misquah  hills,  July  5,  1897,  no.  477.  Tofte  (Carlton  peak), 
July  10,  1897,  nos.  562  and  569.  Tofte,  July  12,  1897,  no. 


Fink  :     LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  243 

649.  Beaver  Bay,  July  13,  1897,  no.  671.  Snowbank  lake 
area,  July  19,  1897,  nos.  806  and  851.  Ely,  July  28,  1897,  no. 
1001. 

17.  Usnea  barbata  (L.)  FR.  var.  ceratina  SCHAER. 

On  trees,  common  or  abundant  but  sterile.  Gunflint,  June 
30,  1897,  no.  263.  Misquah  hills,  July  3,  1897,  no.  423. 
Tofte,  (Carlton  peak),  July  10,  1897,  no.  558.  Beaver  Bay, 
July  13,  1897,  nos.  665  and  672.  Snowbank  lake  area,  July 
19,  1897,  no.  807.  Ely,  July  28,  1897,  no.  993. 

Not  previously  reported  from  Minnesota. 

18.  Usnea  barbata  (L.)  FR.  var.  dasypoga  FR. 

On  trees,  common.     Rose  lake,  June  28,  1897,  no.  213. 

19.  Usnea  barbata  (L.)  FR.  var.  plicata  FR. 

On  trees,  common.  Snowbank  lake  area,  July  19,  1897, 
no.  805.  Ely,  July  28,  1897,  no.  994. 

20.  Usnea  trichodea  ACH. 

On  trees,  common.  Grand  Portage  island,  June  23,  1897, 
no.  157.  English  portage,  June  26,  1897,  no.  190.  Rose  lake, 
June  28,  1897,  no.  212.  Gunflint,  July  2,  1897,  no.  381.  Mis- 
quah Hills,  July  3,  1897,  no.  432,  and  July  5,  1897,  no.  544. 
Tofte  (Carlton  peak),  July  10,  1897,  no.  547.  Tofte,  July  12, 
1897,  no.  659.  Beaver  Bay,  July  13,  1897,  no.  667.  Two 
Harbors,  July  17,  1897,  no.  793. 

Not  previously  reported  from  Minnesota. 

21.  Usnea  longissima  ACH. 

On  trees,  common  along  lake  Superior  and  possibly  in  Mis- 
quah hills.  Sterile.  Grand  Portage  island,  June  18,  1897, 
no.  19.  Misquah  hills,  July  3,  1897,  no.  422.  Tofte,  July 
12,  1897,  no.  649.  Beaver  Bay,  July  13,  1897,  no.  649a. 
Two  Harbors,  July  17,  1897,  no.  787.  Snowbank  lake  area, 
July  20,  1897,  no-  850. 

Specimens  were  collected  at  Grand  Portage  island  five  feet 
long.  The  plant  breaks  with  its  own  weight  and  hangs  abun- 
dantly over  branches  unattached. 

Not  previously  reported  from  Minnesota. 

22.  Usnea  cavernosa  TUCK. 

On  trees,  common.  Grand  Portage  island,  June  17,  1897, 
nos.  6  and  18. 

23.  Alectoria  jubata  (L.)  TUCK. 


244  MINNESOTA    BOTANICAL    STUDIES. 

On  trees  abundant  on  Grand  Portage  island,  infrequent  to 
common  elsewhere.  Sterile.  Grand  Portage  island,  June  18, 
1897,  no.  17.  Gunflint,  July  i,  1897,  no.  354.  Tofte,  July 
12,  1897,  no.  658.  Beaver  Bay,  July  13,  1897,  no.  680. 
Snowbank  lake  area,  July  20,  1897,  no.  837.  Ely,  July  28, 
1897,  no.  996. 

24.  Alectoria  jubata  (L.)  TUCK.  var.  chalybeiformis  ACH. 
On  trees   and  old  wood,   frequent   or   common   throughout. 

Sterile.  High  bluff  at  south  end  of  South  Fowl  lake,  June 
26,  1897,  no.  194.  Gunflint,  June  30,  1897,  no.  258,  and  July 
i,  1897,  no.  316.  Misquah  hills,  July  5,  1897,  no.  514. 
Beaver  Bay,  July  15,  1897,  no.  786.  Snowbank  lake  area, 
July  19,  1897,  no.  815,  and  July  20,  1897,  no.  842.  Ely,  July 
28,  1897,  no.  998. 

25.  Alectoria  jubata  (L.)  TUCK.  var.  implexa  FR. 

On  trees,  infrequent.     Sterile.       Misquah  hills,  July  3,  1897, 
no.  421.     Beaver  Bay,  July  14,  1897,  no.  731. 
Not  previously  reported  from  Minnesota. 

26.  Theloschistes  polycarpus  (EHRH.)  TUCK. 

On  trees,  infrequent  or  rare  west  of  Gunflint,  elsewhere  fre- 
quent or  common.  Grand  Portage  island,  June  18,  1897,  no. 
20.  Gunflint,  July  2,  1897,  no.  393.  Misquah  hills,  July  5, 
1897,  nos.  452  and  487.  Beaver  Bay,  July  13,  1897,  nos.  678 
and  686.  Ely,  July  28,  1897,  no.  985. 

27.  Theloschistes  lychneus  (NYL.)  TUCK. 

On  rocks,  rare.  Grand  Portage  island,  June  23,  1897,  no. 
147.  Snowbank  lake  area,  July  19,  1897,  no.  830. 

28.  Parmelia  perlata  (L.)  ACH. 

On  rocks  and  rarely  on  trees,  rare  to  infrequent  except  at 
Gunflint,  where  the  species  seemed  to  be  common.  Sterile. 
Grand  Portage,  June  24,  1897,  no.  169.  Portage  between 
South  Fowl  lake  and  Pigeon  river,  June  26,  1897,  no.  205. 
Gunflint,  June  7,  1897,  no.  368.  Misquah  hills,  July  5,  1897, 
no.  543.  Beaver  Bay,  July  14,  1897,  no.  725.  Snowbank 
lake  area,  July  24,  1897,  no.  925. 

29.  Parmelia  perforata  (JACQ^)  ACH. 

On  trees,  common.  Sterile.  Tofte  (Carlton  peak),  July 
10,  1897,  no.  572. 

A  puzzling  plant  with  sorediate  margined  lobes  and  otherwise 


Fink  :     LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  245 

resembling  the  last.  However,  the  margins  of  the  lobes  are 
quite  strongly  ciliate,  and  the  lower  surface  of  the  thallus  inter- 
ruptedly so;  possibly  might  be  referred  to  P.  perlata  (L.) 
ACH.  var.  ciliata.  DC.  Thallus  rather  thinner  than  my  her- 
barium specimens  of  P.  perforata. 

30.  Parmelia  perforata  (JACQ^)  ACH.  var.  hypotropa  NYL. 
On  rocks,  rare.     Grand  Portage,  June  23,  1897,  no.  116. 
Not  previously  reported  from  Minnesota. 

31.  Parmelia  crinita  ACH. 

On  rocks  and  once  collected  on  trees,  rare  or  infrequent. 
Sterile.  Grand  Portage,  June  23,  1897,  no.  114.  Gunflint, 
July  i,  1897,  nos.  361  and  362.  Tofte,  July  10,  1897,  no. 
627.  The  plant  differs  from  my  Iowa  and  Ohio  specimens  in 
that  it  is  strongly  ciliate  on  the  upper  surface  of  the  thallus 
among  the  branchlets  and  granules. 

32.  Parmelia  tiliacea  (HOFFM.)  FLOERK. 

On  trees,  rare  or  infrequent.  Gunflint,  July  2,  1897,  no. 
4O7a.  Tofte  (Carlton  peak),  July  10,  1897,  no.  620.  Bea- 
ver Bay,  July  14,  1897,  no.  724a.  Ely,  July  28,  1897,  no. 
1018. 

Really  more  frequent  along  boundary  at  west.  Was  fre- 
quently noted  in  Snowbank  lake  area  and  failure  to  get  speci- 
mens was  an  oversight. 

33.  Parmelia  tiliacea  (HOFFM.)  FLOERK.  var.  sublaevigata  NYL. 
On  trees,  rare.     Snowbank  lake   area,  July  23,   1897,    no. 

896. 

Not  previously  reported  from  Minnesota. 

34.  Parmelia  borreri  TURN.  var.  rudecta  TUCK. 

On  rocks  and  trees,  rare  or  infrequent,  except  common  in  the 
Snowbank  lake  area.  All  sterile  except  no.  744  on  rocks. 
Grand  Portage,  June  24,  1897,  no.  188.  Gunflint,  July  i, 
1897,  no.  369.  Misquah  hills,  July  3,  1897,  no.  431.  Tofte 
(Carlton  peak)  July  10,  1897,  no.  612.  Palisades,  July  15, 
1897,  nos.  744  and  762.  Snowbank  lake  area,  July  30,  1897, 
no.  839.  Ely,  July  28,  1897,  no.  983. 

35.  Parmelia  saxatilis  (L.)  FR. 

On  trees  and  rarely  on  rocks.  Abundant  or  common  and  fre- 
quently fruited.  Grand  Portage  island,  June  17,  1897,  no.  7, 
and  June  21,  1897,  no.  57.  Misquah  hills,  July  3,  1897,  no. 


246  MINNESOTA    BOTANICAL    STUDIES. 

406.     Beaver  Bay,  July  14,  1897,  no.    741.     Snowbank  lake 
area,  July  21,  1897,  no.  873. 

36.  Parmelia  saxatilis  (L.)  FR.  var.  sulcata  NYL. 

On   trees,  probably  common.      Grand   Portage   island,  June 
17,  1897,  no.  7a.     Gunflint,  June  30,  1897,  no.  240. 
Not  previously  reported  from  Minnesota. 

37.  Parmelia  physodes  (L.)  ACH. 

On  trees  and  rocks,  common.  Gunflint,  July  i,  1897,  no. 
383.  Snowbank  lake  area,  July  2,  1897,  no.  883. 

Doubtless  occurring  over  the  whole  region  studied,  but  taken 
for  P.  saxatilis  (L.)  FR.  modified  by  some  peculiarity  of  sub- 
stratum. 

38.  Parmelia  encausta  (SM.)  NYL. 

On  trees,  common.  Grand  Portage  island,  June  6,  1897, 
no.  143.  Tofte  (Carlton  peak),  July  10,  1897,  no.  565. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

39.  Parmelia  olivacea  (L.)  ACH. 

On  trees,  common.  Grand  Portage  island,  June  18,  1897, 
no.  n.  Grand  Portage,  June  23,  1897,  no.  100.  Gunflint,  June 
30,  1897,  no.  260,  and  July  2,  1897,  no.  395.  Tofte  (Carlton 
peak),  July  10,  1897,  nos.  585  and  591.  Beaver  Bay,  July  13, 
1897,  no.  712.  Snowbank  lake  area,  July  20,  1897,  no.  848. 
Ely,  July  28,  1897,  no.  1004. 

40.  Parmelia  olivacea  (L.)  ACH.  var.  prolixa  ACH. 

On  rocks,  common  or  frequent.  High  bluff  at  south  end  of 
South  Fowl  lake,  June  26,  1897,  no.  197.  Gunflint,  June  30, 
1897,  no.  290.  Misquah  hills,  July  5,  1897,  no.  491.  Tofte 
(Carlton  peak),  July  10,  1897,  no.  574.  Beaver  Bay,  July  13, 
1897,  no.  703.  Palisades,  July  15,  1897,  no.  742a.  Snow- 
bank lake  area,  July  20,  1897,  no.  832. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

41.  Parmelia  caperata  (L.)  ACH. 

On  trees  and  rocks,  common.  Grand  Portage  island,  June 
23,  1897,  no.  107.  Gunflint,  June  30,  1897,  no.  254a.  Mis- 
quah hills,  July  3,  1897,  no.  403.  Tofte  (Carlton  peak),  July 
10,  1897,  no  608.  Beaver  Bay,  July  14,  1897,  no.  728.  Pali- 
sades, July  15,  1897,  no.  768.  Snowbank  lake  area,  July  19, 
1897,  no.  804  and  July  24,  1897,  no.  916. 


Fink  :    LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  247 

More  commonly  fruited  than  farther  south  and  especially  well 
fruited  in  the  Snowbank  lake  area. 

42.  Parmelia  conspersa  (EHRH.)  ACH. 

On  rocks,  abundant  or  common.  Grand  Portage  island, 
June  23,  1897,  no.  103.  Gunflint,  June  30,  1897,  no.  289. 
Misquah  hills,  July  5,  1897,  no..  506.  Tofte  (Carlton  peak), 
July  10,  1897,  no.  621.  Palisades,  July  15,  1897,  no.  755. 
Snowbank  lake  area,  July  27,  1897,  no.  967. 

43.  Parmelia  centrifuga  (L.)  ACH. 

On  rocks,  rare  and  sterile.  Misquah  hills,  July  5,  1897, 
no.  496a. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  the  United  States. 

44.  Physcia  speciosa  (WULF.  ACH.)  NYL. 

On  rocks  and  more  rarely  on  old  wood  or  trees,  infrequent  or 
frequent.  Grand  Portage  island,  June  19,  1897,  no.  27.  Gun- 
flint,  June  30,  1897,  no.  270  and  July  i  j  1897,  nos.  342  and  348. 
Misquah  hills,  July  3,  1897,  no.  438,  and  July  5,  1897,  no.  489. 
Tofte  (Carlton  peak),  July  10,  1897,  nos.  571  and  601.  Snow- 
bank lake  area,  July  24,  1897,  no.  911.  Nos.  348,  438  and 
489  fruited. 

45.  Physcia  ciliaris  (L.)  DC. 

On  rocks  or  high  bluffs,  rare.  Grand  Portage,  June  24, 
1897,  no.  180.  Bluffs  at  south  end  of  South  Fowl  lake,  June 
26,  1897,  no.  200.  Palisades,  July  15,  1897,  no.  774. 

Not  previously  reported  from  Minnesota. 

46.  Physcia  aquila  (Acn.)  NYL. 

On  trees,  rare.     Two  Harbors,  July  17,  1897,  no.  791. 
The    only  distinct    specimen    noted    and   not   found   farther 
north. 

Not  previously  reported  from  Minnesota. 

47.  Physcia  pulverulenta  (SCHREB.)  NYL. 

On  rocks  infrequent  or  rare,  and  not  often  fruited.  Grand 
Portage  island,  June  23,  1897,  nos.  145  and  1453.  Grand  Port- 
age, June  23,  1897,  no.  173.  Gunflint,  July  2,  1897?  no.  374. 
Snowbank  lake  area,  July  26,  1897,  no.  964. 

A  very  variable  plant.  No.  1453  is  the  typical  form  with  re- 
spect to  the  upper  surface  of  the  thallus,  being  lighter  colored 
than  the  others  and  pruinose.  Like  the  others  it  is  usually  dark 


248  MINNESOTA    BOTANICAL    STUDIES. 

colored  below.     No.  173  yielded  spores  of  the  usual  size  meas- 
uring   o     -  rnic.  and  much  constricted  in  the  middle.     No.  374 
& 18-23 

gave  spores  only  -  - — -  mic.  and  scarcely  constricted. 

48.  Physcia  pulverulenta  (S^HREB.)  NYL.  var.   leucoleiptes 
TUCK. 

On  rocks  and  trees,  probably  rare.  Sterile.  Grand  Portage 
island»  June  23,  1897,  no.  119.  Misquah  hills,  July  5,  1897, 
no.  496.  Snowbank  lake  area,  July  27,  1897,  no.  966.  Thal- 
lus  black  below. 

Not  previously  reported  from  Minnesota. 

49.  Physcia  stellaris  (L.)  TUCK. 

On  trees  and  rocks,  frequent  or  infrequent.  Grand  Portage 
island,  June  23,  1897,  nos.  105  and  142.  Gunflint,  June  30, 
1897,  no.  283a,  and  July  2,  1897,  no.  394.  Misquah  hills, 
July  5,  1897,  no.  460.  Tofte  (Carlton  peak),  July  10,  1897, 
no.  570.  Beaver  Bay,  July  13,  1897,  no.  700.  Two  Harbors, 
July  17,  1897,  no.  801.  Ely,  July  28,  1897,  no.  1012. 

Occasionally  the  tree  forms  show  dark  fibrils  and  even  dark 
thallus  below  so  that  the  rock  growing  variety,  below,  could 
only  be  distinguished  certainly  by  the  crenulate  border  of  the 
apothecia.  The  plant  is  much  more  variable  than  farther  south 
in  Minnesota  and  Iowa.  No.  570  yielded  apothecia  that  were 
somewhat  dilate  below,  but  the  plant  is  white  below  and  other- 
wise like  the  present  plant  rather  than  P.  obscura  (Ehr.)  Nyl. 

50.  Physcia  stellaris  (L)  TUCK.  var.  apiola  NYL. 

On  rocks,  frequent  or  infrequent.  Grand  Portage  island, 
June  21,  1897,  no.  108.  Gunflint,  June  30,  1897,  no.  284. 
Misquah  hills,  July  5,  1897,  no.  469.  Tofte,  July  12,  1897, 
nos.  64<Da  and  641.  Beaver  Bay,  July  13,  1897,  no.  704. 
Snowbank  lake  area,  July  26,  1897,  no.  953. 

51.  Physcia  tribacia  (AcH.)  TUCK. 

On  rocks,  rare  and  sterile.  Grand  Portage,  June  23,  1897, 
no.  92.  Misquah  hills,  July  5,  1897,  no.  449.  Snowbank 
lake  area,  July  20,  1897,  no.  847. 

52.  Physcia  hispida  (SCHREB.  FR.)  TUCK. 

On  trees,  locally  infrequent  and  poorly  fruited.  Grand  Port- 
age island,  June  21,  1897,  no.  63. 


Fink  :    LICHENS    OF     THE    LAKE    SUPERIOR    REGION.  249 

53.  Physcia  caesia  (HOFFM.)  NYL. 

On  rocks,  frequent  locally.  Sterile.  Gunflint,  June  30, 
1897,  nos.  282  and  292. 

54.  Physcia  obscura  (EHRH.)  NYL. 

On  rocks  and  trees,  rare.  Grand  Portage  island,  June  23, 
1897,  no.  1450.  Grand  Portage,  June  24,  1897,  no.  172. 
Tofte,  July  12,  1897,  no.  642. 

No.  1450  yielded  spores  a  little  large,  -  —  —  mic.,  and  other- 

wise looks  somewhat  like  P.  pulverulent  a  (Schreb.)  Nyl.,  but 
is  black  below.  Apothecia  have  hispid  borders,  and  the  thallus 
is  much  smaller  than  that  of  any  plants  referred  to  the  latter. 

55.  Physcia  adglutinata  (FLOERK.)  NYL. 

On  trees,  rare.  Grand  Portage,  June  23,  1897,  nos.  82a 
and 


56.  Pyxine  sorediata  FR. 

On  rocks  and  trees,  rare  but  widely  distributed.  Collected  in 
fruit  three  times.  Gunflint,  July  I,  1897,  no.  364,  and  July  2, 
1897,  no.  385.  Misquah  hills,  July  3,  1897,  no.  437.  Tofte, 
July  10,  1897,  no.  633.  Palisades,  July  15,  1897,  no.  746. 
Two  Harbors,  July  7,  1897,  no.  790.  Snowbank  lake  area, 
July  24,  1897,  no.  912. 

57.  Umbilicaria  muhlenbergii  (Acn.)  TUCK. 

On  rocks,  common  or  abundant.  Grand  Portage  (Mt.  Jose- 
phine), June  21,  1897,  no.  53.  Gunflint,  June  30,  1897,  nos.  245 
and  246.  Misquah  hills,  July  5,  1897,  no.  537.  Tofte  (Carl- 
ton  peak),  July  10,  1897,  no.  602.  Palisades,  July  15,  1897, 
no.  771.  Snowbank  lake  area,  July  24,  1897,  no.  938. 

58.  Umbilicaria  vellea  (L.)  NYL. 

On  rocks,  common  or  frequent.  Grand  Portage  (Mt.  Jose- 
phine), June  21,  1897,  no.  57,  and  June  24,  1897,  no.  178. 
Gunflint,  July  i,  1897,  no.  367.  Misquah  hills,  July  3,  1897, 
no.  430,  and  July  5,  1897,  no.  538.  Palisades,  July  15,  1897, 
no.  766.  Snowbank  lake  area,  July  24,  1897,  no.  931. 

59.  Umbilicaria  dillenii  TUCK. 

On  rocks,  common.  Grand  Portage,  June  24,  1897,  no.  170. 
Gunflint,  July  I,  1897,  no.  370.  Misquah  hills,  July  3,  1897, 
no.  429,  and  July  5,  1897,  no.  445.  Tofte  (Carlton  peak),  July 


250  MINNESOTA    BOTANICAL     STUDIES. 

10,  1897,  no.  599.  Palisades,  July  15,  1897,  no.  775.  Snow- 
bank lake  area,  July  24,  1897,  no.  937. 

60.  Umbilicaria  pustulata  (L.)  HOFFM.  var.  papulosa  TUCK. 
On  rocks,  rare  to  frequent.     Spores  reaching    100   mic.    in 

length.  Gunflint,  July  i,  1897,  no.  371.  Misquah  hills,  July 
3,  1897,  no.  542.  Palisades,  July  15,  1897,  no.  756.  Snow- 
bank lake  area,  July  24,  1897,  no.  932. 

61.  Umbilicaria  hyperborea  HOFFM. 

On  rocks,  rare,  Grand  Portage  (Mt.  Josephine),  June  23, 
1897,  no.  104. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior within  the  United  States. 

62.  Sticta  amplissima  (Scop.)  MASS. 

On  trees  or  rocks,  rare  or  infrequent.  Grand  Portage,  June 
23,  1897,  no.  115.  Tofte  (Carlton  peak),  July  10,  1897,  no. 
609.  Beaver  Bay,  July  14,  1897,  no.  720.  Two  Harbors. 
July  17,  1897,  no.  802.  Ely,  July  28,  1897,  no.  981. 

63.  Sticta  pulmonaria  (L.)  ACH. 

On  trees  or  rocks,  common  or  frequent.  Grand  Portage 
island,  June  19,  1897,  no.  23.  Rose  lake,  June  28,  1897,  no. 
225.  Gunflint,  July  i,  1897,  no.  365.  Misquah  hills,  July  3, 
1897,  no.  443.  Tofte  (Carlton  peak),  July  10,  1897,  no.  396. 
Beaver  Bay,  July  14,  1897,  no.  724.  Two  Harbors,  July  17, 
1897,  no.  795.  Snowbank  lake  area,  July  19,  1897,  no.  808. 
Ely,  July  28,  1897,  no.  984. 

64.  Sticta  limbata  (SM.)  ACH. 

On  trees,  very  rare.  Tofte,  July  10,  1897,  no.  626.  Only  two 
plants  collected  with  thallus  also  much  like  that  of  the  Euro- 
pean Sticta  fuliginosa.  Spores  brown,  two-celled,  constricted, 


Not  previously  reported  from  Minnesota  and  only  once  from 
North  America.  (Oregon,  by  J.  W.  Eckfeldt.) 

65.  Sticta  crocata  (L.)  ACH. 

On  trees  and  rocks,  rare.  Tofte  (Carlton  peak),  July  10, 
1897,  no  597.  Beaver  Bay,  July  13,  1897,  no.  685.  Snow- 
bank lake  area,  July  24,  1897,  no.  940. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior within  the  United  States. 


Fink  :    LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  251 

66.  Sticta  scorbiculata  (Scop.)  ACH. 

Mossy  rocks  and  trees,  rare  except  at  Grand  Portage  where 
frequent.  Grand  Portage,  June  24,  1897,  no.  176.  Tofte, 
July  10,  1897,  no.  629. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

67.  Nephroma  tomentosum  (HOFFM.)  KOERB. 

On  rocks  and  trees,  frequent.  Grand  Portage  island,  June 
19,  1897,  no.  26.  Misquah  hills,  July  5,  1897,  no.  541.  Tofte, 
July  10,  1897,  no.  634.  Snowbank  lake  area,  July  19,  1897, 
no.  803. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

68.  Nephroma  helveticum  ACH. 

On  rocks  and  occasionally  on  trees  and  earth,  frequent. 
Grand  Portage  island,  June  23,  1897,  no.  133.  Gunflint  June, 
29,  1897,  no.  235,  and  June  30,  1897,  no.  250.  Misquah  hills, 
July  5,  1897,  no.  540.  Beaver  Bay,  July  14,  1897,  no.  740. 
Tofte  (Carlton  peak),  July  10,  1897,  no.  605.  Snowbank 
lake  area,  July  24,  1897,  no.  921,  and  July  26,  1897,  no.  961. 
Ely,  July,  28,  1897,  no.  1005. 

69.  Nephroma  laevigatum  ACH. 

On  trees,  rare.  Misquah  hills,  July  5,  1897,  no.  426.  Two 
Harbors,  July  17,  1897,  no.  799. 

Not  previously  reported  from  Minnesota. 

70.  Nephroma  laevigatum  ACH.  var.  parile  NYL. 

On  rocks,  locally  common.  Grand  Portage,  June  23,  1897, 
no.  113. 

Not  previously  reported  from  Minnesota. 

71.  Peltigera  venosa  (L.)  HOFFM. 

On  earth  and  mossy  rocks,  rare,  spores  reaching  60  mic.  in 
length  and  occasionally  five-celled.  Grand  Portage,  June  23, 
1897,  no.  150.  Portage  at  south  end  of  South  Fowl  lake, 
June  26,  1897,  no.  207. 

72.  Peltigera  aphthosa  (L.)  HOFFM. 

On  earth  or  rocks,  common  or  frequent.  Grand  Portage 
Island,  June  24,  1897,  no.  177.  Gunflint,  June  30,  1897,  no. 
248.  Misquah  hills,  July  5,  1897,  no.  526.  Palisades,  July 
15,  1897,  no.  770.  Snowbank  lake  area,  July  19,  1897,  no.  829. 


252  MINNESOTA    BOTANICAL    STUDIES. 

73.  Peltigera  horizontalis  (L.)  HOFFM. 

On  earth,  frequent  or  common.  Grand  Portage  island,  June 
19,  1897,  no.  30  and  June  23,  1897,  no.  121.  Gunflint,  June 
30,  1897,  no.  247,  and  July  i,  1897,  nos.  338  and  360.  Mis- 
quah  hills,  July  3,  1897,  no.  441.  Tofte,  July  12,  1897,  nos. 
630  and  631.  Beaver  Bay,  July  14,  1897,  no.  723.  Palisades, 
July  15, 1897, no.  761.  Snowbank  lake  area,  July  24,  1897,  nos. 
917  and  918.  Part  of  the  plants  placed  here  agree  somewhat  with 
those  reported  elsewhere  for  Iowa  and  Minnesota  as  P.  •pulveru- 
lenta  (Tayl.)  Nyl.,  but  though  the  sterile  forms  previously  seen 
differ  considerably  from  the  fertile  ones  herein  reported  per- 
haps all  must  eventually  be  placed  here.  The  sterile  forms, 
occasionally  light  colored  below,  are  crisped  and  broken  prob- 
ably from  unfavorable  conditions  which  prevented  their  fruiting. 

Not  previously  reported  from  Minnesota. 

74.  Peltigera  polydactyla  (NECK.)  HOFFM. 

On  earth,  common.  Spores  reaching  no  mic.  Grand  Port- 
age island,  June  23,  1897,  nos.  140  and  144.  Gunflint,  July  i, 
1897,  no.  336.  Misquah  hills,  July  5,  1897,  no.  536.  Beaver 
Bay,  July  14,  1897,  no  726.  Snowbank  lake  area,  July  24, 
1897,  no.  929,  and  July  27,  1897,  no.  971. 

75.  Peltigera  canina  (L.)  HOFFM.  var.  spuria  ACH. 

On  earth,  frequent  or  common.  Grand  Portage  island, 
June  19,  1897,  no.  29.  Grand  Portage  (Mt.  Josephine),  June 
21,  1897,  no.  54.  Gunflint,  July  i,  1897,  no.  323.  Beaver 
Bay,  July  13,  1897,  no.  663.  Palisades,  July  13,  1897,  no. 
760.  Snowbank  lake  area,  July  24,  1897,  no.  943. 

76.  Peltigera  canina  (L.)  HOFFM.  var  sorediata  SCHAER. 

On  earth,  frequent.  Grand  Portage  (Mt.  Josephine),  June 
23,  1897  no.  u8a.  Gunflint,  July  i,  1897,  no.  343.  Mis- 
quah hills,  July  5,  1897,  no.  535.  Tofte  (Carlton  peak),  July 
10,  1897,  no.  589.  Beaver  Bay,  July  13,  1897,  no.  693. 
Snowbank  lake  area,  July  24,  1897,  no.  902.  Ely,  July  28, 
1897,  no.  976. 

77.  Solorina  saccata  (L.)  ACH. 

On  earth,  rare.  Grand  Portage  island,  June  24,  1897,  no. 
179.  Not  previously  reported  from  Minnesota  and  new  to  the 
interior  of  North  America. 

78.  Pannaria  languinosa  (Acn.)  KOERB. 

On  rocks,  common.      Grand  Portage  island,  June  17,  i8( 


Fink  :    LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  253 

no.  i.  South  Fowl  lake,  June  26,  1897,  no.  195.  Gunflint, 
June  30,  1897,  no.  272.  Misquah  hills,  July  3,  1897,  no.  435, 
and  July  5,  1897,  no.  457.  Tofte  (Carlton  peak),  July  10, 
1897,  no.  56ia.  Beaver  Bay,  July  13,  1897,  no.  691.  Snow- 
bank lake  area,  July  19,  1897,  no.  824.  Nos.  195  and  457 
showing  the  bright  sulphur-colored  plant  common  in  Europe 
and  only  noted  in  North  America  by  the  present  writer  in  the 
second  paper  of  this  series. 

79.  Pannaria  microphylla  (Sw.)  DELIS. 

On  rocks,  frequent  on  Grand  Portage  island,  June  19,  1897, 
no.  22.  Gunflint,  July  i,  1897,  no.  330.  Beaver  Bay,  July 
13,  1897,  no.  684.  Ely,  July  28,  1897,  no.  987. 

80.  Pannaria  lepidiota  TH.  FR. 

On  rocks  and  wood,  infrequent.  Grand  Portage,  June  24, 
1897,  no.  175.  Gunflint,  July  2,  1897,  no.  372.  Misquah 
hills,  July  5,  1897,  nos.  463  and  479.  Tofte  (Carlton  peak), 
July  10,  1897,  no.  582.  Snowbank  lake  area,  July  24,  1897, 
no.  908. 

Not  previously  reported  from  Minnesota. 

81.  Pannaria  flabellosa  TUCK. 

On  rocks,  rare.  Grand  Portage  island,  June  23,  1897,  no. 
128.  Sterile,  but  having  the  narrow  linear  lobed  thallus  and 
blue-black  hypothallus.  The  thallus  does  not  show  the  ex- 
panded and  striated  lobes  at  circumference. 

Not  previously  reported  from  Minnesota  and  new  west  of  New 
England. 

82.  Pannaria  nigra  (HUDS.)  NYL,. 

On  rocks,  rare.      Gunflint,  July  i,  1897,  no.  347. 

83.  Ephebe  pubescens  FR. 

On  rocks,  rare,  sterile.  Palisades,  July  13,  1897,  no.  745. 
Snowbank  lake  area,  July  24,  1897,  no.  901. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

84.  Ephebe  solida  BORN.  (?) 

On  rocks,  rare  and  sterile.  Misquah  hills,  July  5,  1897,  no. 
488.  Beaver  Bay,  July  13,  1897,  no.  683.  Snowbank  lake 
area,  July  20,  1897,  no.  849. 

A  short  form  growing  in  small  dense  tufts. 


254  MINNESOTA    BOTANICAL    STUDIES. 

85.  Collema  pycnocarpum  NYL. 

On  trees,  rare.  Snowbank  lake  area,  July  27,  1897,  nos. 
968  and  974. 

86.  Collema  flaccidum  ACH.  (?) 

On  high  rocks,  rare.  Misquah  hills,  July  5,  1897,  no.  495. 
Sterile,  but  with  the  thallus  corresponding  with  tree  forms 
farther  south,  except  that  the  plant  is  larger. 

87.  Collema  nigrescens  (Huos.)  ACH. 

On  trees,  especially  Populus,  common  or  frequent.  Grand 
Portage,  June  19,  1897,  no.  36,  and  June  23,  1897,  no.  101. 
Rose  lake,  June  28,  1897,  no.  221.  Gunflint,  July  30,  1897, 
no.  266.  Misquah  hills,  July  5,  1897,  no.  483.  Beaver  Bay, 
July  15,  1897,  no.  783.  Snowbank  lake  area,  July  19,  1897,  no. 
822,  and  July  21,  1879,  no.  874.  Ely,  July  28,  1897,  no.  1014. 

88.  Collema f urvum  ( ACH.)  NYL? 

On  wet  rocks,   frequent.      Grand   Portage  island,    June  23, 
1897,  no.  156.     Tofte,  July  12,  1897,  no.  646.     Not  typical. 
Not  previously  reported  from  Minnesota. 

89.  Leptogium  lacerum  (Sw.)  FR. 

On  rocks,  rare  and  sterile.  Snowbank  lake  area,  July  21, 
1897,  no.  867. 

90.  Leptogium  lacerum  (Sw.)  FR.  var.  pulvinatum  MOUG.  and 
NESTL. 

On  rocks,  rare.  Snowbank  lake  area,  July  29,  1897,  no.  965. 
Not  previously  reported  from  Minnesota. 

91.  Leptogium  tremelloides  (L.)  FR. 

On  rocks  and  rarely  on  trees.  Widely  distributed,  but  usually 
rare  locally.  Portage  between  South  Fowl  lake  and  Pigeon 
river,  June  26,  1897,  no.  208.  Gunflint,  July  I,  1897,  nos. 
349  and  358.  Misquah  hills,  July  3,  1897,  no.  424.  Beaver 
Bay,  July  15,  1897,  no.  781.  Snowbank  lake  area,  July  20, 
1897,  no.  843,  July  26,  1897,  no.  962,  and  July  27,  1897,  no- 
974a.  Ely,  July  28,  1897,  nos.  977  and  1008. 

92.  Leptogium  myochroum  (EHRH.,  SCHA.ER.)  TUCK. 

On  trees  and  rocks,  frequent.  Grand  Portage  island,  June 
19,  1897,  no.  24.  Gunflint,  July  i,  1897,  no.  341.  Tofte 
(Carlton  peak),  July  10,  1897,  no.  598.  Snowbank  lake  area 
July  19,  1897,  no.  820. 

Not  previously  reported  from  Minnesota. 


Fink:    LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  255 

93.  Leptogium    myochroum    (£HRH.,    SCHAER.)    TUCK.    var. 
tomentosum  SCHAER. 

On  trees,  rare.     Tofte  (Carlton  peak),  July  10,  1897,  no.  559. 
Not  previously  reported  from  Minnesota. 

94.  Placodium  elegans  (LINK.)  DC. 

^  On  rocks,  common.  Grand  Portage,  June  21,  1897,  no.  69. 
Grand  Portage  island,  June  23,  1897,  nos.  93  and  98.  Gun- 
flint,  June  30,  1897,  no.  294,  and  July  i,  1897,  no.  328.  Mis- 
quah  hills,  July  5,  1897,  no.  444.  Palisades,  July  15,  1897, 
no.  747.  Snowbank  lake  area,  July  22,  1897,  no.  887,  no. 
444,  looking  toward  the  next  in  having  orange-red  apothecia. 

95.  Placodium  murorum  (HOFFM.)  DC. 

On  rocks,  rare.      Misquah  hills,  July  5,  1897,  no.  450. 
Not  previously  reported  from  Minnesota. 

96.  Placodium  murorum  (HOFFM.)  DC.  var.  miniatum  TUCK. 
On  rocks,  rare.   Sterile.  Grand  Portage,  June  23,  1897,  no.  88. 
Not  previously  reported  from  Minnesota. 

97.  Placodium  cinnabarinum  (Acn.)  ANZ. 

On  rocks,  common  at  the  one  locality.  Gunflint,  July  2 
no.  378. 

98.  Placodium  citrinura  (HOFFM.)  LEIGHT. 

On  rocks,  infrequent.  No.  68  well  fruited  but  with  thallus 
nearly  obsolete  in  some  specimens.  Grand  Portage  island, 
June  21,  1897,  no.  68.  Grand  Portage,  June  23,  1897,  no.  117. 
Misquah  hills,  July  5,  1897,  no.  461. 

99.  Placodium  aurantiacum  (LIGHTF.)  NAEG.  and  HEPP. 

On  rocks,  rare  at  first  locality,  frequent  at  second.  Grand 
Portage  island,  June  23,  1897,  no.  127.  Gunflint,  June  30, 
1897.  nos.  296  and  298,  and  July  2,  1897,  no.  389.  Beaver 
Bay,  July  13,  1897,  no.  661.  No.  296  with  a  white  thallus  and 
)therwise  not  typical,  but  I  can  place  it  nowhere  else. 

100.  Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP. 

On  trees,  frequent.     Grand   Portage,  June  23,  1897,  no.  84. 
Gunflint,  June   30,  1897,  no.  253,  and  July   2,  1897,   no.  392. 
leaver  Bay,  July  13,  1897,  no.  669. 

101.  Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP.  var.  pyra- 
cea  NYL. 

On  old  wood,  common  locally.     Beaver  Bay,  July  13,  1807, 
no.  682. 


256  MINNESOTA    BOTANICAL    STUDIES. 

102.  Placodium  vitellinum  (EHRH.)  NAEG.  and  HEPP. 

On  old  wood,  frequent.  Beaver  Bay,  July  13,  1897,  no.  660. 
Spores  simple  or  two-celled  and  reaching  30  in  each  ascus. 

103.  Placodium   vitellinum  (EHRH.)    NAEG.  and  HEPP.    var. 
aurellum  ACH. 

On  rocks,  common  at  Gunflint,  rare  elsewhere.  Grand  Port- 
age island,  June  24,  1897,  no.  163.  Gunflint,  June  30,  1897, 
no.  279,  and  July  i,  1897,  no.  329.  Tofte  (Carleton  peak), 
July  10,  1897,  no.  618.  Snowbank  lake  area,  July  24,  1897, 
no.  903.  Spores  reaching  20  in  asci. 

104.  Lecanora  rubina  (VILL.)  ACH. 

On  rocks,  common  or  frequent.  Grand  Portage  (Mt.  Jose- 
phine), June  19,  1897,  no.  47.  Grand  Portage  island,  June  21, 
1897,  no.  73.  Gunflint,  June  30,  1897,  no.  265.  Misquah 
hills,  July  5,  1897,  no.  486.  Beaver  Bay,  July  13,  1897,  no. 
705.  Snowbank  lake  area,  July  26,  1897,  no.  054. 

105.  Lecanora  rubina  (VILL.)  ACH.  var.  heteromorpha  ACH. 
On  rocks,  frequent  locally.     Grand  Portage  (Mt.  Josephine), 

June  19,  1897,  no.  47a.     Gunflint,  July  i,  1897,  no.  326.     Pali- 
sades, July  15,  1897,  no.  748. 

Not  previously  reported  from  Minnesota. 

106.  Lecanora     muralis    (SCHREB.)     SCHAER.     var.     saxicola 

SCHAER. 

On  rocks,  common  or  frequent.  Grand  Portage  island,  June 
21,  1897,  no.  75.  Rose  lake,  June  29,  1897,  no.  224.  Mis- 
quah Chills,  July  5,  1897,  no.  513.  Tofte,  July  12,  1897,  no.  643. 

107.  Lecanora  muralis  (SCHREB.)  SCHAER.  var.  diffracta  FR. 

On  rocks,  rare.  Grand  Portage  (Mt.  Josephine),  June  19, 
1897,  no.  46. 

Not  previously  reported  from  Minnesota. 

108.  Lecanora  pallida  (SCHREB.)  SCHAER. 

On  trees,  infrequent.     Gunflint,  July  i,  1897,  no.  337. 
Not  previously  reported  from  Minnesota. 

109.  Lecanora  frustulosa  (DICKS.)  MASS. 

On  rocks,  frequent.  Grand  Portage,  June  21,  1897,  no.  78, 
and  June  23,  1897,  no.  153.  Gunflint,  July  2,  1897,  no.  376. 
Beaver  Bay,  July,  13,  1897,  no.  706. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 


Fink  :    LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  257 

1 10.  Lecanora  sordida  (PERS.)  TH.  FR. 

Rocks,  common  at  the  last  two  locations  which  were  high 
bluffs.  Grand  Portage,  June  24,  1897,  no.  185.  South  Fowl 
lake,  June  26,  1897,  no.  198.  Misquah  hills,  July  5,  1897, 
nos.  465  and  505. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

in.  Lecanora  subfusca  (L.)  ACH. 

On  trees  and  rocks,  common  on  the  former.  Grand  Portage, 
June  23,  1897,  no.  131.  South  end  of  South  Fowl  lake,  June 
26,  1897,  no.  202.  Misquah  hills,  July  5,  1897,  no.  500. 
Gunflint,  June  30,  1897,  no.  274,  and  July  i,  1897,  no.  305. 
Tofte  (Carlton  peak),  July  10,  1897,  nos.  588  and  595.  Beaver 
Bay,  July  15,  1897,  no.  780.  Snowbank  lake  area,  July  19, 
1897,  no.  825,  and  July  21,  1897,  no.  855. 

112.  Lecanora  subfusca  (L.)  ACH.  var.  hypnorum  SCHAER. 
Among  moss  on  a  cedar  tree,  rare.     Misquah  hills,  July  5, 

1897,  no.  494a.     Not  previously  reported  from  Minnesota. 

113.  Lecanora  subfusca  (L.)  ACH.  var.  coilocarpa  Ach. 

On  trees  and  rocks,  frequent.  Grand  Portage  island,  June 
21,  1897,  no.  67.  Gunflint,  June  30,  1897,  no.  301,  and  July 
i,  1897,  no.  339.  Misquah  hills,  July  5,  1897,  no.  481. 
Beaver  Bay,  July  13,  1897,  no.  698. 

114.  Lecanora  varia  (EHRH.)  NYL. 

On  rocks,  rare  or  infrequent  and  some  specimens  perhaps 
approaching  var.  -polytropa  Nyl.  Grand  Portage,  June  19, 
1897,  no.  41.  Gunflint,  July  i,  1897,  no.  307,  and  July  2, 
1897,  no.  380. 

115.  Lecanora  varia  (EHRH.)  NYL.  var.  saepincola  FR. 

On  wood,  common 'at  first  locality.     Beaver  Bay,  July  14, 
1897,  no.   742.     Snowbank  lake  area,  July  22,  1897,  no.  889. 
Not  previously  reported  from  Minnesota. 

116.  Lecanora  varia  (EHRH.)  NYL.  var.  symmicta  ACH. 

On  old  wood,  rare.  Grand  Portage  island,  June  18,  1897, 
no.  12.  Gunflint,  July  2,  1897,  no.  400.  Tofte  (Carlton  peak), 
July  10,  1897,  no.  580. 

117.  Lecanora  hageni  ACH. 

On  rocks,  rare.     Gunflint,  July  i,  1897,  no.  357- 

118.  Lecanora  hageni  ACH.  var.  sambuci  (PERS.)  TUCK. 


258  MINNESOTA    BOTANICAL    STUDIES. 

On  trees  common  locally.  Grand  Portage  island,  June  19, 
1897,  no.  25.  Misquah  hills,  July  5,  1897,  no.  501.  Tofte 
(Carlton  peak),  July  10,  1897,  no.  575. 

A  puzzling  plant  with  exciple  commonly  entire  or  excluded 
and  looking  quite  as  much  like  forms  of  L.  subfusca  (L.)  Ach. 
or  L.  varia.  (Ehrh.)  Nyl.  Spores  reaching  sixteen  in  asci. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

119.  Lecanora  elatina  ACH. 

On  trees,  rare.     Tofte,  July  10,  1897,  no.  638. 
Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

120.  Lecanora  pallescens  (L.)  SCHAER. 

On  trees,  infrequent  or  rare,  but  widely  distributed.  Mis- 
quah hills,  July  3,  1897,  no.  412,  and  July  5,  1897,  no.  494. 
Tofte  (Carlton  peak),  July  10,  1897,  no.  592.  Beaver  Bay, 
July  13,  1897,  no.  674.  Two  Harbors,  July  17,  1897,  no.  798. 
Snowbank  lake  area,  July  21,  1897,  no.  864,  and  July  27, 
1897,  no.  973. 

Not  previously  reported  from  Minnesota. 

121.  Lecanora  tartarea  (L.)  ACH. 

On  rocks,  rare.     Ely,  July  28,  1897,  no.  988. 
Not  previously  reported  from  Minnesota. 

122.  Lecanora  cinerea  (L.)  SOMMERF. 

On  rocks,  common  or  abundant.  The  thallus  varying  in 
color  from  ash-color  to  a  dull  black.  Grand  Portage  (Mt. 
Josephine),  June  19,  1897,  no.  39.  Grand  Portage  island, 
June  21,  1897,  no.  74.  Gunflint,  June  30,  1897,  nos.  285,  293a, 
295  and  300.  Misquah  hills,  July  5,  1897,  nos.  458,  468  and 
4933.  Tofte  (Carlton  peak),  July  10,  1897,  nos.  615  and  622. 
Palisades,  July  15,  1897,  no.  752.  Beaver  Bay,  July  15,  1897, 
no.  778.  Snowbank  lake  area,  July  20,  1897,  no.  833.  Ely, 
July  28,  1897,  no.  990. 

123.  Lecanora  cinerea  (L.)  SOMMERF.  var.  laevata  FR. 

On  rocks,  rare.  Grand  Portage,  June  23,  1897,  no.  94. 
Gunflint,  June  30,  1897,  no.  286.  Misquah  hills,  July  5,  1897, 
no.  471.  Snowbank  lake  area,  July  20,  1897,  no.  836. 

124.  Lecanora  cinerea  (L.)  SOMMERF.  var.  gibbosa  NYL. 

On  rocks,  rather  rare.  Grand  Portage  island,  June  23,  1897, 
no.  79.  Gunflint,  July  i,  1897,  no.  3093.  Misquah  hills,  July 
5,  1897,  no.  472. 


Fink  :    LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  259 

125.  Lecanora  calcarea  (L.)  SOMMERF.  (?) 

On  rocks,  rare.     Thallus   almost  obsolete   and    spores    only 

— *  mic.     Beaver  Bay,  July  13,  1897,  no.  707. 
5-7 
Not  previously  reported  from  Minnesota. 

126.  Lecanora  calcarea  (L.)  SOMMERF.  var.  contorta  FR. 

On  rocks,  rare.      Grand  Portage  island,  June  23,  1897,  no. 

155. 

Not  previously  reported  from  Minnesota. 

127.  Lecanora  fuscata  (SCHRAD.)  TH.  FR. 

On  rocks,  frequent.  Grand  Portage  (Mt.  Josephine),  June 
19,  1897,  no.  44a.  Grand  Portage,  June  21,  1897,  no.  71. 
Gunflint,  July  i,  1897,  no.  331. 

128.  Lecanora  fuscata  (SCHRAD.)  Tn.  FR.  var.  rufescens  TH. 
FR. 

On  rocks,  frequent  at  second  locality.  Misquah  hills,  July 
5,  1897,  no.  446.  Beaver  Bay,  July  14,  1897,  no.  718,  and  July 
15,  1897,  no.  777. 

Not  previously  reported  from  Minnesota. 

129.  Rinodina  oreina  (AcH.)  MASS. 

On  rocks,  rare,  preferring  high  perpendicular  rocks  or  larger 
masses  of  talus.  No.  750  is  an  unusually  coarse  form,  but  must 
be  referred  here.  South  end  of  South  Fowl  lake,  June  26, 
1897,  no.  203.  Misquah  hills,  July  5,  1897,  no.  518.  Pali- 
sades, July  15,  1897,  nos.  750  and  753.  Snowbank  lake  area 
[uly  20,  1897,  no.  854. 

130.  Rinodina  ascociscana  TUCK. 

On    trees,    rare.     Gunflint,   July    i,  1897,    no.    340.     Tofte 
(Carlton  peak),  July  10,  1897,  no.  594. 
Not  previously  reported  from  Minnesota. 

131.  Rinodina  sophodes  (AcH.)  NYL. 

Abundant  on  drift  pebbles  at  Beaver  Bay,  infrequent  on  wood 
elsewhere.  Grand  Portage,  June  23,  1897,  no.  95.  Tofte, 
[uly  12,  1897,  no.  651.  Beaver  Bay,  July  13,  1897,  no.  702. 
Snowbank  lake  area,  July  20,  1897,  no.  853. 

132.  Rinodina  sophodes  (AcH.)  NYL.  var.  confragosa  NYL. 

On  wood,  rare.  Spores  25  to  34  mic.  in  length.  Snowbank 
lake  area,  July  21,  1897,  no.  875. 

Not  previously  reported  from  Minnesota. 


260  MINNESOTA    BOTANICAL    STUDIES. 

133.  Pertusaria  velata  (TURN.)  NYL. 

On  trees,  infrequent.  Portage,  between  Rose  and  Rove 
lakes,  June  27,  1897,  no.  2ioa.  Tofte  (Carlton  peak),  July 
10,  1897,  no.  577.  Ely,  July  28,  1897,  no.  1017. 

134.  Pertusaria  multipuncta  (TURN.)  NYL. 

On  trees,  frequent  at  second  locality.  Tofte  (Carlton  peak), 
July  10,  1897,  no.  611.  Snowbank  lake  area,  July  28,  1897, 
no.  900. 

135.  Pertusaria  multipunctata  (TURN.)   NYL.   var.   laevigata 
TURN,  and  BORR. 

On  trees,  probably  frequent.  Grand  Portage,  June  23,  1897, 
no.  158.  Rose  lake,  June  28,  1897,  no.  2i7a.  Gunflint,  July 
2,  1897,  no.  389a.  Misquah  hills,  July  5,  1897,  no.  478. 

Not  previously  reported  from  Minnesota  and  new  to  North 
America. 

136.  Pertusaria  communis  DC. 

On  trees,  nearly  always  cedars,  common  or  frequent.  Port- 
age, between  Rose  and  Rove  lakes,  June  27,  1897,  no.  210. 
Rose  lake,  June  28,  1897,  no.  217.  Gunflint,  July  i,  1897, 
no.  336,  Misquah  hills,  July  7,  1897,  nos.  453  and  499. 
Beaver  Bay,  July  13,  1897,  no.  664,  and  July  15,  1897,  no. 
782.  Snowbank  lake  area,  July  21,  1897,  nos.  870  and  888. 

137.  Pertusaria  sp. 

On  trees.  Spores  nearly  like  the  above  in  the  few  apothecia 
not  transformed  into  soredia.  Thallus  lighter  colored  at  cir- 
cumference with  frequent  two  or  three  dark  lines  near  circum- 
ference. Misquah  hills,  July  3,  1897,  no.  410.  Beaver  Bay, 
July  15,  1897,  no.  688.  Snowbank  lake  area,  July  19,  1897, 
no.  826. 

138.  Pertusaria  leioplaca  (Acn.)  SCHAER. 

On  trees,  widely  distributed  but  seldom  common  in  any  local- 
ity. Grand  Portage,  June  23,  1897,  no.  91.  Portage  between 
North  Fowl  lake  and  Moose  lake,  June  26,  1897,  no.  192. 
Rose  lake,  June  28,  1897,  no.  218.  Gunflint,  June  30,  1897, 
no.  275,  and  July  2,  1897,  no.  399.  Misquah  hills,  July  5, 
1897,  no.  491  a.  Beaver  Bay,  July  15,  1897,  no.  784.  Ely, 
July  28,  1897,  no.  1010.  Varying  greatly  according  to  sub- 
stratum. On  young  trees  with  smooth  bark  the  thallus  is  thin 
and  smooth  and  the  ostioles  frequently  indistinct.  On  older 
trees  with  rough  bark  the  thallus  is  thicker  and  broken,  and 


Fink  :  LICHENS  OF  THE  LAKE   SUPERIOR  REGION.  261 

the  apothecia  are  falsely  lecanoroid.  The  last  feature  is  due 
no  doubt  to  great  age  of  these  plants  which  began  growth  when 
the  trees  were  young.  The  extremes  appear  macroscopically 
like  distinct  species. 

Not  previously  reported  from  Minnesota. 

139.  Pertusaria  pustulata  (Acn.)  NYL. 

On  trees,  infrequent.  Rose  lake,  June  28,  1897,  no.  228. 
Gunflint,  July  i,  1897,  no.  310.  Misquah  hills,  July  3,  1897, 
no.  418. 

140.  Pertusaria  glomerata  (Acn.)  SCHAER. 

On  rocks,  very  rare.     Misquah  hills,  July  5,  1897,  no.   490. 
Not  previously  reported  from   Minnesota  and  new  to  the  in- 
terior of  North  America. 

141.  Gyalecta  fagicola  (HEPP.)  TUCK. 

On  trees,  rare.  Snowbank  lake  area,  July  22,  1897,  no. 
880. 

Not  previously  reported  from  Minnesota  and  new  to  west  of 
of  New  England. 

142.  Urceolaria  scruposa  (L.)  NYL. 

On  rocks,  rare  or  infrequent,  but  widely  distributed  and  no. 
126  with  thallus  approaching  var.  gypsacea  Nyl.  Grand  Port- 
age island,  June  25,  1897,  no.  126.  Gunflint,  June  30,  1897, 
no.  302.  Misquah  hills,  July  5,  1897,  no.  473.  Beaver  Bay, 
July  13,  1897,  no.  701.  Snowbank  lake  area,  July  23,  1897, 
no.  899.  Ely,  July  28,  1897,  no.  978. 

143.  Stereocaulon  coralloides  FR. 

On  rocks,  frequent.     Tofte  (Carlton  peak),  July  10,  1897.  no. 

549- 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

144.  Stereocaulon  paschale  (L.)  FR. 

On  earth  among  rocks,  common.  Grand  Portage  (Mt. 
Josephine),  June  19,  1897,  no.  49.  Grand  Portage  island, 
June  21,  1897,  no.  50.  Gunflint,  June  30,  1897,  no.  233. 
Misquah  hills,  July  5,  1897,110.  482.  Beaver  Bay,  July  13, 
1897,  no.  689.  Palisades,  July  15,  1897,  no.  758.  Snowbank 
lake  area,  July  24,  1897,  no.  927.  No.  49  seems  to  approach 
the  above.  Also  no.  50  has  the  stout  podetia  of  S.  tomentosum 
(Fr.)  Th.  Fr.  and  is  somewhat  tomentose.  Yet  it  appears 
nearer  herb,  specimens  of  the  above. 


262  MINNESOTA    BOTANICAL    STUDIES. 

145.  Cladonia  symphycarpa  FR.  var.  epiphylla  (Acn.)  NYL. 
In  crevices  in  rocks,  rare.   Habitat  unusual  but  thallus  too  large 

for  C.  ccesptticia  (Pers.)  Fl.    Gunflint,  July  I,  1897,  no.  363. 
Not  previously  reported  from  Minnesota. 

146.  Cladonia  mitrula  TUCK. 

On  earth,  rare.     Beaver  Bay,  July  15,  1897,  no.  694. 

147.  Cladonia  cariosa  (Acn.)  SPRENG. 

On  earth,  probably  frequent.  Grand  Portage  island,  June 
23,  1897,  nos.  120  and  146.  Gunflint,  July  i,  1897,  no.  324. 
Misquah  hills,  July  5,  1897,  nos.  509  and  533.  Tofte  (Carl- 
ton  peak),  July  10,  1897,  no.  606.  Beaver  Bay,  July  14,  1897, 
nos.  737  and  739. 

The  forms  listed  here  seem  to  me  to  be  partly  intermediate 
between  this  and  the  last  having  the  habit  of  this,  but  some  are 
rather  small  with  the  squamules  usually  small.  C.  mitrula  Tuck, 
is  the  common  form  in  southern  Minnesota,  but  the  better  de- 
velopment of  this  region  runs  into  the  present  species.  Of  the 
specimens  here  listed  no.  146  is  the  best  representative  of  the 
species  and  no.  120  the  poorest.  Some  of  the  smaller  ap- 
proaches C.  symphycarpa  Fr.,  which  is  itself  a  doubtful  species. 

148.  Cladonia  decorticata  FLOERK. 

On  earth,  rare.  Beaver  Bay,  July  14,  1897,  no.  738.  Not 
previously  reported  from  Minnesota  and  new  to  west  of  New 
England. 

149.  Cladonia  pyxidata  (L.)  FR. 

On  earth,  common.  Grand  Portage  island,  June  19,  1897, 
no.  32.  Gunflint,  July  i,  1897,  nos.  333  and  352.  Misquah 
hills,  July  5,  1897,  no.  534.  Tofte  (Carlton  peak),  July  10, 
1897,  nos.  567  and  568.  Beaver  Bay,  July  14,  1897,  no.  735. 
Snowbank  lake  area,  July  21,  1897,  no.  868,  and  July  27, 
1897,  no.  972. 

150.  Cladonia  fimbriata  (L.)  FR. 

On  earth,  rare.      Gunflint,  June  30,  1897,  no.  259. 
Not  previously  reported  from  Minnesota. 

151.  Cladonia  fimbriata  (L.)  FR.  var.  tubaeformis  FR. 

On  dead  wood  and  earth,  common.  Grand  Portage  island, 
June  23,  1897,  no.  129.  Gunflint,  June  30,  1897,  no.  268. 
Misquah  hills,  July  3,  1897,  no.  439.  Tofte  (Carlton  peak), 
July  10,  1897,  no.  561.  Snowbank  lake  area,  July  24,  1897, 
no.  930. 


Fink:    LICHENS    OF     THK    LAKE    SUPERIOR    REGION.  2G3 

152.  Cladonia  fimbriata  (L.)  Fr.  var.  radiata  Fr. 

On  earth  and  old  wood,  frequent.  Grand  Portage  island, 
June  19,  1897,  no.  33  and  June  24,  1897,  no.  162.  Gunflint, 
June  30,  1897,  no.  234.  Snowbank  lake  area,  July  20,  1897, 
no.  840. 

Not  previously  reported  from  Minnesota. 

153.  Cladonia  gracilis  (L.)  NYL. 

On  earth,  common  or  abundant  and  extremely  variable. 
Grand  Portage  island,  June  19,  1897,  no.  35.  Gunflint,  July 
i,  1897,  nos.  325,  344  and  345.  Misquah  hills,  July  5,  1897, 
nos.  508  and  522.  Tofte  (Carlton  peak),  July  10,  1897,  nos. 
550,  564  and  604.  Beaver  Bay,  July  14,  1897,  no.  734.  Snow- 
bank lake  area,  July  21,  1897,  no  876  and  July  24,  1897,  nos. 
922  and  923. 

154.  Cladonia  gracilis  (L.)  NYL.  var.  verticillata  FR. 

On  earth,  rare,  Grand  Portage  island,  June  19,  1897,  nos. 
35b  and  35d.  Tofte  (Carlton  peak),  July  10,  1897,  no.  625. 

155.  Cladonia  gracilis  (L.)  NYL.  var.  symphycarpia  TUCK. 
On  earth,  infrequent,  possibly  as  near  C.  degenerans  Floerk. 

Grand  Portage  island,  June  19,  1897,  no.  35a. 
Not  previously  reported  from  Minnesota. 

156.  Cladonia  gracilis  (L.)  NYL.  var.  cervicornis  FLOERK. 
On  earth,  rare.     Gunflint,  June  30,  1897,  no.  231. 

Not  previously  reported  from  Minnesota. 

157.  Cladonia  gracilis  (L.)  NYL.  var.  hybrida  SCHAER. 

On  earth,  common.  Grand  Portage  island,  June  19,  1897, 
no.  35c.  Gunflint,  July  I,  1897^0.346.  Misquah  hills,  July 
3,  1897,  no.  433.  Snowbank  lake  area,  July  19,  1897,  no. 
821,  and  July  24,  1897,  no.  905. 

158.  Cladonia  turgida  (£HRH.)  HOFFM. 

On  earth,  common  at  Gunflint.     Gunflint,  June  30,  1897,  nos. 
241  and  252.     Snowbank  lake  area,  July  24,  1897,  no.  928. 
Not  previously  reported  from  Minnesota. 

159.  Cladonia  turgida  (£HRH.)  HOFFM. var.  conspicua  (SCHAER.) 
NYL. 

On  earth,  frequent.  Rose  lake,  June  28,  1897,  no.  237. 
Misquah  hills,  July  3,  1897,  no.  425,  and  July  5,  1897,  no. 
525.  Tofte,  (Carlton  peak),  July  10,  1897,  nos.  603  and  637. 

Not  previously  reported  from  Minnesota. 


264  MINNESOTA    BOTANICAL    STUDIES. 

1 60.  Cladonia  squamosa  HOFFM. 

On  earth,  common  or  abundant.  Grand  Portage  island, 
June  24,  1897,  no.  165.  Gunflint,  June  30,  1897,  no.  232,  and 
July  i,  1897,  no.  350,  and  July  2,  1897,  no.  384.  Misquah 
hills,  July  5,  1897,  no.  529.  Tofte,  July  12,  1897,  nos.  628 
and  632.  Beaver  Bay,  July  14,  1897,  no.  719.  Above  Pali- 
sades, July  15,  1897,  no.  772.  Snowbank  lake  area,  July  24, 
1897,  nos.  913,  919  and  924.  Ely,  July  28,  1897,  no.  995. 

161.  Cladonia  squamosa  HOFFM.  var.  phyllocoma  RABENH. 
On   earth,  frequent.      Grand  Portage  island,  June  23,  1897, 

no.  141.     Misquah    hills,  July  5,  1897,   no.    459.     Snowbank 
lake  area,  July  24,  1897,  nos.  939  and  942. 

Not  previously  reported  from  Minnesota  and  new  to  North 
America. 

162.  Cladonia  cornuta  (L.)  FR. 

On   earth,  rare.     Grand  Portage  island,  June  24,  1897,  no. 
159.     Misquah  hills,  July  5,  1897,  no.  511. 
Not  previously  reported  from  Minnesota. 

163.  Cladonia  delicata  (EHRH.)  FL. 

On  old  wood,  rare.     Beaver  Bay,  July  13,  1897,  no.  692. 

164.  Cladonia  caespiticia  (PERS.)  FL. 

On  old  wood,  rare.  Tofte  (Carlton  peak),  July  10,  1897, 
no.  586. 

165.  Cladonia  furcata  (Huos.)  FR. 

On  earth,  frequent.  Gunflint,  June  30,  1897,  nos.  236  and 
237.  Misquah  hills,  July  5,  1897,  no.  524.  Above  Palisades, 
July  15,  1897,  no,  767.  Snowbank  lake  area,  July  24,  1897, 
no.  914. 

166.  Cladonia  furcata  (Huos.)  FR.  var.  crispata  FL. 

On  earth,  common  locally.  Grand  Portage,  June  24,  1897, 
no.  168.  Gunflint,  June  30,  1897,  no.  249.  Palisades,  July 
15,  1897,  no.  757. 

167.  Cladonia  rangiferina  (L.)  HOFFM. 

On  earth,  abundant  or  common.  Grand  Portage  island, 
June  17,  1897,  no.  2,  and  June  23,  1897,  135.  Grand  Portage 
(Mt.  Josephine),  June  19,  1897,  no.  48.  Gunflint,  June  30, 
1897,  no.  244.  Misquah  hills,  July  5,  1897,  no.  521.  Tofte, 
(Carlton  peak),  July  10,  1897,  no.  548.  Beaver  Bay,  July  14, 
1897,  no.  732.  Palisades,  July  15,  1897,  no.  759.  Snowbank 
lake  area,  July  24,  1897,  no.  926. 


Fink  :    LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  265 

1 68.  Cladonia  rangiferina  (L.)  HOFFM.  var.  sylvatica  L. 

On  earth,  frequent.  Misquah  hills,  July  5,  1897,  no.  531. 
Tofte  (Carlton  peak),  July  10,  1897,  no.  553.  Snowbank  lake 
area,  July  24,  1897,  nos.  915  and  936. 

169.  Cladonia  rangiferina  (L.)  HOFFM.  var.  alpestris  L. 

On  earth,  common  locally.  Grand  Portage,  June  24,  1897, 
no.  171.  Gunflint,  July  i,  1897,  no.  351.  Above  Palisades, 
July  15,  1897,  no.  773. 

170.  Cladonia  amaurocraea  (FL.)  SCHAER. 

On  earth  and  rocks,  common  or  frequent.  Grand  Portage 
(Mt.  Josephine),  June  21,  1897,  no.  52.  Grand  Portage  island, 
June  24,  1897,  no.  164.  Gunflint,  June  30,  1897,  no.  243. 
Misquah  hills,  July,  5,  1897,  nos.  530  and  532.  Tofte  (Carl- 
ton  peak),  July,  10,  1897,  no.  552.  Palisades,  July  15,  1897, 
no.  769.  Snowbank  lake  area,  July  24,  1897,  nos.  935  and 
941. 

Not  previously  reported  from  Minnesota. 

171.  Cladonia  uncialis  (L.)  FR. 

On  earth,  frequent  or  common.  Grand  Portage  (Mt.  Jose- 
phine), June  21,  1897 ,'no.  55.  Grand  Portage,  June  24,  1897, 
no.  167.  Gunflint,  June  30,  1897,  no.  239.  Misquah  hills,  July 
5,1897,110.475.  Misquah  hills,  July  5,  1897,  no.  528.  Tofte 
(Carlton  peak),  July  10,  1897,  no.  551.  Beaver  Bay,  July  14, 
1897,  no.  727.  Palisades,  July  15,  1897,  no.  764.  Snowbank 
lake  area,  July  24,  1897,  no.  933. 

172.  Cladonia  cornucopioides  (L.)  FR. 

On  earth,  rare  but  widely  distributed.  Grand  Portage,  June 
24,  1897,  no.  166.  Gunflint,  July  30,  1897,  no.  238.  Misquah 
hills,  July  5,  1897,  nos.  497  and  527.  Tofte  (Carlton  peak), 
July  10,  1897,  no.  563.  Above  Palisades,  July  15,  1897,  no. 

754- 

173.  Cladonia  deformis  (L.)  HOFFM. 

On  earth,  rare.     Grand  Portage  island,  June  23,  1897,  no. 

IS1- 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

174.  Cladonia  digitata  (L.)  HOFFM. 

On  an  old  stump,  rare.     Tofte,  July  12,  1897,  no.  655. 
Not  previously  reported  from  Minnesota  and   new  to  the  in- 
terior of  North  America. 


266  MINNESOTA    BOTANICAL    STUDIES. 

175.  Cladonia  macilenta  (EHRH.)  HOFFM. 

On  old  wood  and  earth,  rare.  Misquah  hills,  July  5,  1897, 
no.  510.  Above  Palisades,  July  15,  1897,  no.  776. 

176.  Cladonia  cristatella  TUCK. 

On  earth  and  old  wood,  abundant.  Grand  Portage  island, 
June  18,  1897,  no.  31.  Gunflint,  July  i,  1897,  nos.  322  and  332. 
Misquah  hills,  July  3,  1897,  no.  417.  Tofte  (Carlton  peak), 
July  10,  1897,  no.  610.  Beaver  Bay,  July  14,  1897,  no.  736. 
Snowbank  lake  area,  July  19,  1897,  no.  827. 

177.  Baeomyces  byssoides  (L.)  SCHAER. 

On  rocks,  rare.   Grand  Portage  island,  June  21,  1897,  no.  109. 
Not  previously  reported  from  Minnesota,  and  new  to  the  in- 
terior of  North  America. 

178.  Baeomyces  seruginosus  (Scop.)  DC. 

On  rotton  wood,  common.  Grand  Portage  island,  June  18, 
1897,  no.  16.  Gunflint,  July  i,  1897,  no.  309.  Misquah  hills, 
July  3,  1897,  no.  404.  Tofte,  July  12,  1897,  no.  639.  Beaver 
Bay,  July  14,  1897,  no.  721.  Snowbank  lake  area,  July  19, 
1897,  no.  823. 

Not  previously  reported  from  Minnesota,  and  new  to  the  in- 
terior of  North  America. 

179.  Biatora  rufonigra  TUCK. 

On  rocks,  frequent.  Grand  Portage  (Mt.  Josephine),  June 
19,  1897,  no.  37.  Grand  Portage  island,  June  21,  1897,  no. 
64.  South  Fowl  lake,  June  26,  1897,  no.  193.  Misquah  hills, 
July  5,  1897,  no.  456.  Snowbank  lake  area,  July  21,  1897, 
no.  859. 

180.  Biatora  coarctata  (SM.,  NYL.)  TUCK. 

On  rocks,  rare.     Beaver  Bay,  July  13,  1897,  no.  709. 
Not  previously  reported  from  Minnesota. 

181.  Biatora  viridescens  (SCHRAD.)  FR. 

On  old  wood,  common  locally.  Misquah  hills,  July  3,  1897, 
no.  414. 

Snowbank  lake  area,  July  20,  1897,  no.  846. 

Not  previously  reported  from  Minnesota,  and  new  to  the  in- 
terior of  North  America. 

182.  Biatora  vernalis  (L.)  FR. 

On  old  wood,  mosses  and  trees,  frequent.     Grand  Portage 


LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  267 

island,  June   21,    1897,   no.   61,   and  June  23,    1897,   no.    149. 
Gunnint,  June  30,  1897,  no.  262. 

Not  previously  reported  from  Minnesota. 

183.  Biatora  sanguineoatra  (FR.)  TUCK. 

On  earth  common.  Grand  Portage  island,  June  19,  1897, 
no.  21  and  28.  Grand  Portage,  June  24,  1897,  no.  181.  Gun- 
flint,  June  30,  1897,  no.  269,  and  July  i,  1897,  no.  359.  Mis- 
quah  hills,  July  5,  1897,  no.  413.  Tofte  (Carlton  peak),  July 
10,  1897,  no.  578.  Snowbank  lake  area,  July  22,  1897,  no.  884. 

Not  previously  reported  from  Minnesota. 

184.  Biatora  turgidula  (FR.)  NYL. 

On  old  wood,  rare.  Grand  Portage  island,  June  18,  1897, 
no.  14. 

Not  previously  reported  from  Minnesota. 

185.  Biatora  leucophaea  (FI.OERK). 

On  rocks,  infrequent.      Grand  Portage,  June  23,    1897,   no. 
139,  and  June  24,  1897,  no.  187.    Gunflint,  July  i,  1897,  no.  306. 
Not  previously  reported  from  Minnesota. 

186.  Biatora  leucophaea  FLOERK  var.  griseoatra  KOERB. 
On  rocks,  rare.     Grand  Portage,  June  24,  1897,  no.  186. 
Not  previously  reported  from  Minnesota  and  new  south  of 

Arctic  America. 

187.  Biatora  uliginosa  (SCHRAD.)  FR. 

On  earth,  abundant  near  Disappointment  lake.  Beaver  Bay, 
July  13,  1897,  no.  708.  Snowbank  lake  area,  July  24,  1897, 
no.  944.  Ely,  July  28,  1897,  no.  997. 

Not  previously  reported  from  Minnesota. 

188.  Biatora  sphaeroides  (DICKS.)  TUCK. 

On  old  wood,  rare.  Snowbank  lake  area,  July  24,  1897,  no. 
944a. 

Not  previously  reported  from  Minnesota. 

189.  Biatora  glauconigrans  TUCK. 

On  trees,  rare.     Gunnint,  July  i,  1897,  no.  398. 
Not  previously  reported    from    Minnesota  and  new  west  of 
New  England. 

190.  Biatora  arthropurpurea  (MASS.)  HEPP. 

On  trees,  rare.  Grand  Portage  island,  June  21,  1897,  no. 
56.  Snowbank  lake  area,  July  22,  1897,  no.  811.  Ely,  July 
28,  1897,  no.  1007. 


268  MINNESOTA    BOTANICAL    STUDIES. 

191.  Biatora  oxyspora  (TuL.)  NYL. 

On  Parmelia  colpodes,  rare.  Misquah  hills,  July  3,  1897, 
no.  416. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

192.  Biatora  lucida  (Acn.)  FR. 

On  damp  rocks,  rare.  Grand  Portage,  June  23,  1887,  no. 
183. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

193.  Biatora  myriocarpoides  (NYL.)  TUCK. 

On  old  wood,  frequent  locally.  Beaver  Bay,  July  13,  1897, 
no.  673.  Apothecia  larger  than  usual. 

194.  Biatora  flavidolivens  TUCK. 

On  old  wood,  frequent  locally.  Rose  lake,  June  28,  1897, 
no.  214.  Misquah  hills,  July  5,  1897,  no.  504. 

Not  previously  reported  from  Minnesota  and  new  west  of  New 
England.  • 

195.  Biatora  hypnophila  (TURN.)  TUCK. 

On  trees,  rare.  Gunflint,  June  30,  1897,  no.  273.  Snow- 
bank lake  area,  July  26,  1897,  no.  956. 

196.  Biatora  nsegelii  HEPP. 

On  old  wood,  rare.  Beaver  Bay,  July  13,  1897,  no.  715. 
Not  previously  reported  from  Minnesota  and  new  west  of  New 
England. 

197.  Biatora  rubella  (EHRH.)  RABENH. 

On  trees,  rare  or  infrequent.  Tofte  (Carlton  peak),  July  ior 
1897,  no.  546.  Beaver  Bay,  July  13,  1897,  no.  640.  Snow- 
bank lake  area,  July  19,  1897,  no.  812,  July  21,  1897,  no.  872, 
and  July  24,  1897,  no.  906.  Ely,  July  28,  1897,  no.  1021. 

198.  Biatora  fuscorubella  (HOFFM.)  TUCK. 

On  trees,  infrequent.  Snowbank  lake  area,  July  17,  1897, 
no.  816,  and  July  21,  1897,  no.  861. 

199.  Biatora  schweinitzii  FR. 

On  cedars,  rare.     Misquah  hills,  July  5,  1897,  no.  493. 
Not  previously  reported  from  Minnesota. 

200.  Biatora  incompta  (BORR.)  HEPP. 

On    trees,  probably  common  locally.      Rose   lake,  June  28, 
1897,  no.  222.     Gunflint,  July  i,  1897,  no.  335. 
Not  previously  reported  from  Minnesota. 


Fink  :    LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  269 

201.  Biatora  muscorum  (Sw.)  TUCK. 

On  trees  with  moss,  rare.  Snowbank  lake  area,  July  26, 
1897,  no.  963. 

202.  Heterothecium  sanguinarium  (L.)  FLOT. 

On  old  wood  and  occasionally  on  trees  or  rocks,  common  ex- 
cept in  last  region.  Gunflint,  July  i,  1897,  nos.  318  and  319. 
Misquah  hills,  July  3,  1897,  nos.  409  and  415,  and  July  5, 
1897,  nos.  492a  and  498.  Tofte  (Carlton  peak),  July  10,  1897, 
no.  581.  Beaver  Bay,  July  15,  1897,  no.  695.  Two  Harbors, 
July  17,  1897,  no.  796.  Snowbank  lake  area,  July  19,  1897, 
no.  810. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

203.  Heterothecium  sanguinarium  (L.)  FLOT.  var.  affine  TUCK. 
On  wood,  rare.     Rose  lake,  June   28,  1897,  no.  222a. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

204.  Lecidea  lactea  FL. 

On  rocks  along  lake  Superior,  common  especially  north. 
Grand  Portage  island,  June  21,  1897,  no.  76,  and  June  23, 
1897,  no.  136.  Grand  Portage,  June  23,  1897,  no.  182.  Tofte, 
July  12,  1897,  no.  652.  Palisades,  July  15,  1897,  no.  751. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

205.  Lecidea  crustulata  ACH. 

On  rocks,  rare.     Grand   Portage  island,  June  23,  1897,   no. 

137- 

Not  previously  reported  from  Minnesota  and  known  else- 
where in  North  America  only  from  Labrador  by  Eckfeldt  and 
Arnold. 

206.  Lecidea  lapicida  FR. 

On  rocks,  probably  common  locally.  Misquah  hills,  July  5, 
1897,  no.  448. 

Not  previously  reported  from  Minnesota  and  new  to  the 
interior  of  North  America. 

207.  Lecidea  lapicida  FR.  var.  oxydata  FR. 

On   rocks,    rare.     Grand   Portage,  June  24,    1897,  no.   174. 
Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 


270  MINNESOTA    BOTANICAL    STUDIES. 

208.  Lecidea  speirea  NYL. 

On  rocks  along  lake  shore,  rare.  Grand  Portage  island, 
June  23,  1897,  no.  no. 

Not  previously  reported  from  Minnesota. 

209.  Lecidea  albocaerulescens  (WULF.)  SCHAER. 

On  rocks,  rare.     Misquah  hills,  July  5,  1897,  no.  484. 

210.  Lecidea  platycarpa  ACH. 

On  rocks,  rare.     Misquah  hills,  July  5,  1897,  no.  474. 
Not  previously  reported  from  Minnesota. 

211.  Lecidea  enteroleuca  FR. 

On  trees  and  rocks,  common.  Grand  Portage  island,  June 
21,  1897,  nos.  59b  and  60,  and  June  23,  1897,  no.  in.  Grand 
Portage,  June  23,  1897,  no.  148.  English  Portage,  June  26, 
1897,  no.  191.  Misquah  hills,  July  5,  1897,  no.  492. 
Beaver  Bay,  July  13,  1897,  no.  700.  Snowbank  lake  area, 
July  24,  1897,  no.  909. 

212.  Lecidea  enteroleuca  FR.  var.  achrista  SOMMERF. 

On  trees,  infrequent.     Grand  Portage  island,  June  21,  1897, 
no.    59a.     Grand  Portage,  June    23,    1897,  no.  97. 
Not  previously  reported  from  Minnesota. 

213.  Lecidea  melancheima  TUCK. 

On  old  wood,  rare.  Gunflint,  July  2,  1897,  no.  390.  Mis- 
quah hills,  July  5,  1897,  no.  454.  Snowbank  lake  area,  July 
20,  1897,  no.  835. 

Not  previously  reported  from  Minnesota. 

214.  Lecidea  cyrtidia  TUCK. 

On  pebbles,  rare,  thallus  reduced  and  hence  the  black  hypo- 
thallus  prominent.     Snowbank  lake  area,  July  27,  1897,  no.  969. 
Not  previously  reported  from  Minnesota. 

215.  Lecidea  acclinis  FLOT. 

On  cedars,  rare.     Gunflint,  July  i,  1897,  no.  353. 
Not  previously  reported  from  Minnesota. 

216.  Buellia  alboatra  (HOFFM.)  TH.  FR. 
On  rocks,  rare. 

Grand  Portage,  June  23,  1897,  no.  99. 
Not  previously  reported  from  Minnesota. 

217.  Buellia  parasema  (AcH.)  TH.  FR. 

On  trees,  common.     Ely  specimen  having  spores  reaching 

mic.     Grand  Portage  island,  June  16,  1897,  no.  4,  and 


Fink:    LICHENS    OF     THE    LAKE    SUPERIOR    REGION.  271 

June  21,  1897,  no.  65.  South  Fowl  lake,  June  26,  1897,  no. 
196.  Gunflint,  July  i,  1897,  nos.  311  and  312.  Misquah  hills, 
July  5,  1897,  no.  480.  Beaver  Bay,  July  15,  1897,  no.  676. 
Snowbank  lake  area,  July  19,  1897,  nos.  8na,  813  and  817. 
Ely,  July  28,  1897,  no.  989a. 

218.  Buellia  parasema  (Acn.)  TH.  FR.  var.  triphragmia  NYL. 
On  trees,  infrequent.     Gunflint,  June  30,  1897,  nos.  255  and 

278.     Tofte,  July  10,  1897,  no.  613. 

Not  previously  reported  from  Minnesota. 

219.  Buellia  dialyta  (NYL.)  TUCK. 

On  pines,  rare.     Two  Harbors,  July~~i7,  1897,  no.  792. 
Not  previously  reported  from  Minnesota,  and  new  to  the  in- 
terior of  North  America. 

220.  Buellia  myriocarpa  (DC.)  MUDD. 

On  old  wood,  abundant  locally.  Beaver  Bay,  July  14,  1897, 
no.  7i6a. 

221.  Buellia  myriocarpa  (DC.)  MUDD.  var.  polyspora  WILLEY. 
On  trees,  rare.     Ely,  July  28,  1897,  no.  ion. 

222.  Buellia  petraea   (FLOT.,  KOERB.)  TUCK. 

On  rocks,  common  or  abundant.  Grand  Portage  island, 
June  17,  1897,  no.  5.  Gunflint,  June  30,  1897,  nos.  281,  283 
and  291.  Misquah  hills,  July  5,  1897,  no.  470.  Beaver  Bay, 
July  15,  1897,  no.  779.  Ely,  July  28,  1897,  no.  989. 

223.  Buellia  petraea    (FLOT.,   KOERB.)    TUCK.    var.  grandis 
FLOERK. 

On  rocks,  rare.  Gunflint,  June  30,  1897,  no.  293,  and  July 
i,  1897,  no.  304. 

224.  Buellia  petraei  (FLOT.,  KOERB.)  TUCK.  var.  montagnaei 
TUCK. 

On  rocks,  common  or  abundant.  Grand  Portage  (Mt.  Jose- 
phine), June  18,  1897,  no.  42.  Gunflint,  July  2,  1897,  no.  379. 
Misquah  hills,  July  5,  1897,  no.  467.  Beaver  Bay,  July  13, 
1897,  no.  662.  Palisades,  July  15,  1897,  no.  749.  Snowbank 
lake  area,  July  20,  1897,  no.  834. 

225.  Buellia  geographica  (L. )  TUCK. 

On  rocks,  rare  and  approaching  var.  lecanorina  Flcerk 
Gunflint,  June  30,  1897,  no.  303.  Palisades,  July  15,  1897,  no. 

743- 

Not  previously  reported  from  Minnesota,  and  new  to  the  in- 
terior of  North  America. 


272  MINNESOTA    BOTANICAL     STUDIES. 

226.  Buellia  parmeliarum  (SOMMERF.)  TUCK. 

On  Parmelia  borreri,  rare.  Snowbank  lake  area,  July  22, 
1897,  no.  885. 

Not  previously  reported  from  Minnesota. 

227.  Opegrapha  varia  (PERS.)  FR. 

On  trees,  common  on  cedars  except  along  lake  Superior. 
Gunflint,  July  i,  1897,  no.  334.  Misquah  hills,  July  5,  1897, 
no.  502.  Tofte  (Carlton  peak),  July  10,  1897,  no.  593.  Snow- 
bank lake  area,  July  21,  1897,  no.  857. 

228.  Opegrapha  varia  (PERS.)  TUCK.  FR.  var.  notha  ACH. 
On  cedars,  locally  abundant.       Rose  lake,  June  28,   1897, 

no.  220. 

Not  previously  reported  from  Minnesota. 

229.  Graphis  scripta  (L.)  ACH. 

On  trees,  frequent  or  common.  Grand  Portage  island,  June 
21,  1897,  no.  66.  Gunflint,  July  i,  1897,  nos.  314  and  317. 
Tofte  (Carlton  peak),  July  10,  1897,  no.  576.  Beaver  Bay, 
July  15,  1897,  no.  716.  Snowbank  lake  area,  July  21,  1897, 
no.  858. 

230.  Graphis  scripta  (L.)  ACH.  var.  recta  (HUMB.)  NYL. 

On  birch  trees,  infrequent.  Grand  Portage,  June  24,  1897, 
no.  189.  Misquah  hills,  July  3,  1897,  no.  405.  Tofte,  July 
12,  1897,  no.  650.  Snowbank  lake  area,  July  23,  1897,  no. 
897. 

231.  Graphis  scripta  (L.)  ACH.  var.  limitata  ACH. 

On  trees,  very  rare.      Resembles    G.  dendritica  externally  as 
to  apothecia.     Misquah  hills,  July  5,  1897,  no.  451. 
Not  previously  reported  from  Minnesota. 

232.  Arthonia  dispersa  (SCHRAD.)  NYL. 

On  Acer  spicatum,  abundant.  Grand  Portage,  June  23,  1897, 
no.  86.  Tofte  (Carlton  peak),  July  10,  1897,  no.  590.  Beaver 
Bay,  July  13,  1897,  no.  714.  Snowbank  lake  area,  July  20, 
1897,  no.  845.  Ely,  July  28,  1897,  no.  979. 

Not  previously  reported  from  Minnesota. 

233.  Arthonia  radiata  (PERS.)  TH.  FR. 

On  trees  in  low  places,  common.  Grand  Portage  island, 
June  23,  1897,  no.  123.  Rose  lake,  June  28,  1897,  no.  219. 
Gunflint,  July  i,  1897,  no.  315,  and  July  2,  1897,  no.  388. 
Tofte  (Carlton  peak),  July  10,  1897,  no.  584,  Beaver  Bay, 


Fink  :    LICHENS    OF    THE    LAKE    SUPERIOR    REGION.  273 

July  15,  1897,  no.  785.     Snowbank  lake  area,  July  20,  1897, 
no.  838.     Ely,  July  28,  1897,  no.  1013. 

234.  Arthonia  punctiformis  ACH. 

On  trees,  locally  common.  Gunflint,  June  30,  1897,  no.  277. 
Misquah  hills,  July  3,  1897,  no.  407. 

235.  Arthonia  patellulata  NYL. 

On  trees,  rare.      Gunflint,  June  30,  1897,  no.  254. 
Not  previously  reported  from  Minnesota. 

236.  Calicium  trichiale  ACH. 

On  trees,  common  locally.  Rose  lake,  June  28,  1897,  no. 
230.  Beaver  Bay,  July  15,  1897,  no.  696.  Snowbank  lake 
area,  July  19,  1897,  no.  818. 

Not  previously  reported  from  Minnesota,  and  new  to  the  in- 
terior of  North  America. 

237.  Calicium  trichiale  ACH.  var.  stemoneum  NYL. 
On  pine,  common.      Ely,  July  28,  1897,  no.  992. 

Not  previously  reported  from  Minnesota,  and  new  to  the  in- 
terior of  North  America. 

238.  Calicium  brunneolum  ACH. 

On  decorticated  wood,  common  locally.  Two  Harbors,  July 
17,  1897,  no.  800.  Snowbank  lake  area,  July  12,  1897,  no. 
860.  Ely,  July  26,  1897,  no.  1000. 

Not  previously  reported  from  Minnesota,  and  new  to  the  in- 
terior of  North  America. 

239.  Calicium  chrysocephalum  (TURN.)  ACH. 

On  trees,  frequent.  Misquah  hills,  July  5,  1897,  no.  447. 
Two  Harbors,  July  17,  1897,  no. -788.  Snowbank  lake  area, 
July  22,  1897,  no.  882.  Ely,  July  28,  1897,  no.  1003. 

Not  previously  reported  from  Minnesota,  and  new  to  the  in- 
terior of  North  America. 

240.  Calicium  chrysocephalum  (TURN.)  ACH.  var.  filare  SCH. 
On  cedars,  rare.     Tofte,  July  12,  1897,  no.  647. 

Not  previously  reported  from  Minnesota.  Variety  apparently 
new  to  North  America. 

241.  Calicium  parietinum  ACH. 

On  decorticated  wood,  common.  Grand  Portage  island,  June 
17,  1897,  no.  3.  Gunflint,  July  2,  1897,  no.  382.  Misquah 
hills,  July  3,  1897,  no.  408. 

Not  previously  reported  from  Minnesota. 


274  MINNESOTA    BOTANICAL    STUDIES. 

242.  Calicium  quercinum  PERS. 

On   dead   wood,  infrequent.     Rose   lake,  June  28,  1897,  no. 
229.     Tofte  (Carlton  peak),  July  10,  1897,  no  583. 
Not  previously  reported  from  Minnesota. 

243.  Calicium  hyprellum  ACH.  var.  viride  NYL. 

On  trees,  rare.  Misquah  hills,  July  5,  1897,  no.  476.  Snow- 
bank lake  area,  July  21,  1897,  no.  877,  and  July  22,  1897,  no. 
893.  Ely,  July  28,  1897,  no.  1015. 

Not  previously  reported  from  Minnesota.  Variety  new  to 
North  America.  Stipes  sometimes  very  short. 

244.  Calicium  turbinatum  PERS. 

On  Pertusaria  communis,  rare.     Beaver  Bay,  July  13,  1897, 
no.  6643.     Snowbank  lake  area,  July  21,  1897,  no.  866. 
Not  previously  reported  from  Minnesota. 

245.  Coniocybe  pallida  (PERS.)  FR. 

On  Fraxinus,  rare.  Snowbank  lake  area,  July  19,  1897, 
no.  831. 

Not  previously  reported  from  Minnesota. 

246.  Endocarpon  miniatum  (L.)  SCHAER. 

On  rocks,  along  shore  of  lake  Superior,  very  rare.  Grand 
Portage  island,  June  21,  1897,  no.  80. 

247.  Endocarpon    miniatum  (L.)  SCHAER.   var.    complicatum 
SCHAER. 

On  rocks,  frequently  1000  feet  above  water  level,  frequent. 
Grand  Portage  (Mt.  Josephine),  June  19,  1897,110.  38.  Grand 
Portage  island,  June  23,  1897,  no  102.  Misquah  hills,  July  5> 
1897,  no.  445. 

248.  Endocarpon  fluviatile  DC. 

On  rocks  frequently  inundated,  common.  Rose  lake,  June 
28,  1897,  no.  211.  Gunflint,  July  i,  1897,  no.  327.  Misquah 
hills,  July  7,  1897,  no.  512.  Snowbank  lake  area,  July  21, 
1897,  no.  878. 

249.  Thelocarpon  prasinellum  NYL. 

On  rocks,  rare.  Grand  Portage  (Mt.  Josephine),  June  22, 
1897,  no.  90. 

The  plant  agrees  here  and  not  with  saxicoline  species,  Euro- 
pean or  American. 

250.  Staurothele  umbrina  (WAHL.)  TUCK. 

Wet    rocks,     common.     Misquah   hills,  July   5,    1897,   nos. 


Fink  :     LICHENS    OF    THE    LAKE    SUPERIOR    REGION  275 

462,  519   and   520.     Snowbank   lake   area,  July  26,  1897,  no. 

957- 

Not  previously  reported  from  Minnesota. 

251.  Staurothele  drummondii  TUCK. 

On  rocks  along  the  shore,  frequent  locally.  Grand  Portage 
island,  June  21,  1897,  no.  72. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

252.  Verrucaria  nigrescens  PERS. 

On  rocks,  rare.     Grand  Portage,  June  24,  1897,  no.  184. 

253.  Verrucaria  epigaea  (PERS.)  ACH. 

On  earth,   rare.     Snowbank   lake   area,  July  26,   1897,  no. 

944- 

Not  previously  reported  from  Minnesota. 

254.  Sagedia  oxyspora  (NvL.)  TUCK. 

On   birch,  rare.     Beaver  Bay,  July  13,  1897,  no.  697. 
Not  previously  reported   from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

255.  Pyrenula  punctiformis  (Acn.)  NAEG.  var.  fallax  NYL. 
On  trees,  common.     Gunflint,  June  30,  1897,  no.  276.    Mis- 

quah  hills,  July  5,  1897,  no.  503.     Snowbank   lake   area,  July 
21,  1897,  no.  871,  and  July  26,  1897,  no.  950. 
Not  previously  reported  from  Minnesota. 

256.  Pyrenula  leucoplaca  (WALLR.)  KRB. 

On  trees,  common  to  west  of  region.  Between  Rose  and 
Rove  lakes,  June  27,  1897,  no.  209.  Gunflint,  July  2,  1897, 
no.  391.  Misquah  hills,  July  5,  1897,  nos.  499  and  507. 
Snowbank  lake  area,  July  19,  1897,  no.  828,  July  20,  1897,  no. 
852,  and  July  26,  1897,  nos.  955,  958  and  959.  Ely,  July  28, 
1897,  nos.  1009  and  1019. 

257.  Pyrenula  cinerella  (FLOT.)  TUCK. 

On  birch,  common.  Grand  Portage  island,  June  21,  1897, 
no.  56. 

The  only  American  specimens  seen  by  me  which  show  the 
spores  as  large  as  those  of  the  European  plant.  Spores  meas- 
ured 12-18  by  6-9  mic.  Spore  measurements  for  the  species  in 
America  are  more  commonly  12—17  by  5-7  mic.,  my  Iowa 
specimens  giving  12-16  by  6-7  mic.,  and  T.  A.  William's  from 
Nebraska,  15-17  by  5^2-7  mic. 

Not  previously  reported  from  Minnesota. 


276  MINNESOTA    BOTANICAL    STUDIES. 

258.  Pyrenula  cinerella  (PLOT.)  TUCK.  var.  quadriloculata,  var. 

nov. 

Spores  12-15  by  5-6^  mic.,  passing  from  2  and  occasion- 
ally 3-celled  to  a  much  more  common  4-celled  condition.  The 
apothecia  somewhat  below  normal  size  for  the  species.  Pyren- 
ula punctiformis  Ach.,  Naeg.  vw.fallax  Nyl.,  quite  commonly 
occurs  with  the  species  and  variety,  as  it  does  with  the  latter  in 
the  present  instance  and  with  the  former  both  in  Minnesota  and 
Iowa. 

On  birch,  probably  common  locally.  Grand  Portage  island, 
June  24,  1897,  no.  85. 


XIX.     CONTRIBUTIONS     TO    A     KNOWLEDGE    OF 

THE   LICHENS    OF   MINNESOTA.— V.  LICHENS 

OF   THE    MINNESOTA   VALLEY    AND 

SOUTHWESTERN  MINNESOTA. 


BRUCE  FINK. 


CONSIDERATIONS   OF   DISTRIBUTION    AND    HABITAT. 

The  area  considered  in  this  paper  was  selected  with  a  view 
to  obtaining  as  complete  a  knowledge  as  possible  of  the  lichen 
flora  of  the  Minnesota  river  valley  and  of  that  of  southwestern 
Minnesota  in  general. 

The  upper  portion  of  the  valley  near  Minneapolis  would,  of 
course,  give  a  flora  essentially  like  that  of  Minneapolis  and  vi- 
cinity already  studied.  Hence,  for  the  month's  field  work,  it 
was  thought  best  to  begin  operations  at  a  locality  a  considerable 
distance  from  Minneapolis.  As  an  initial  place,  Mankato,  about 
60  miles  from  Minneapolis,  was  selected.  The  location  of  this 
city  is  also  advantageous  in  that  it  lies  nearly  midway  between 
the  Minneapolis  and  the  northeastern  Iowa  areas  compared 
carefully  in  the  second  paper  of  this  series,  thus  forming  a  con- 
necting link  between  the  two  areas  previously  studied.  After 
a  careful  study  of  the  lichens  of  the  Mankato  area  both  to  gain 
a  knowledge  of  the  lichen  flora  of  the  region  and  for  the  sake 
of  relationships  with  the  areas  indicated  above,  New  Ulm  was 
next  selected  as  an  area  of  special  interest  because  of  the  expo- 
sures of  Cretaceous  sandstone  and  the  most  southeastward  ex- 
posures of  quartzite  rocks  in  the  valley.  At  New  Ulm  only 
these  two  rock  formations  were  studied,  as  time  spent  on  other 
substrata  present  would  only  be  repaid  for  most  part  by  a  repe- 
tition of  the  species  found  upon  the  same  substrata  at  Mankato, 
only  30  miles  distant.  Three  days  were  next  spent  at  Red- 
wood Falls,  Morton  and  North  Redwood  with  a  view  to  secur- 
ing rare  species  and  noting  the  southeastern  extension  of  certain 
species  in  the  valley.  From  here  I  proceeded  to  Granite  Falls. 

277 


278  MINNESOTA    BOTANICAL    STUDIES. 

This  being  the  most  northwestern  area  reached  in  the  survey,  its 
lichen  flora  was  studied  carefully.  The  final  task  was  to  study 
the  lichen  flora  of  the  pipestone  and  the  Sioux  quartzite  at 
Pipestone. 

A  brief  statement  as  to  substrata  is  next  in  order.  About 
Mankato  trees  abound,  and  three  kinds  of  rock — limestone, 
sandstone  and  bowlders — are  plentiful.  I  found  only  the  two 
interesting  substrata  mentioned  above  atNewUlm.  Trees  and 
bowlders  were  abundant,  but  were  not  studied  for  the  reason  al- 
ready stated.  At  Redwood  Falls,  Morton  and  North  Redwood, 
granite  trees  and  earth  were  examined  for  species  especially 
rare  or  interesting.  The  great  masses  of  granite,  supposed  to 
have  been  exposed  since  the  close  of  the  glacial  age,  formed  the 
most  interesting  substratum  at  Granite  Falls.  This  is  also  the 
most  northwestern  area  in  the  valley  where  trees  occur  in  any 
considerable  numbers.  The  calcareous  drift  pebbles  and  cal- 
careous earth  proved  also  very  interesting  here.  The  two  sub- 
strata examined  at  Pipestone  have  been  mentioned.  I  need  to 
add  only  one  statement  more  to  make  the  analysis  of  substrata 
complete  enough  for  the  present  purpose.  This  is  that  earth 
was  examined  everywhere  and  furnished  much  of  interest,  as 
will  appear  later. 

The  following  rare  lichens  were  found  only  at  Redwood 
Falls,  Morton  or  North  Redwood :  Peltigera  canina  (L.) 
Hoffm.  var.  spongiosa  Tuck,  and  Stereocaulon  -paschale  (L.) 
Fr.  Also  the  area  including  the  above  places  forms  the  most 
southeastern  known  extension  of  the  following  lichens  in  the 
valley:  Parmelia  olivacea  (L.)  Ach.  vw.prolixa  Ach.  ;  Pan- 
naria  microphylla  (Sw.)  Delis;  Omphalaria  phyllisca  (Wahl.) 
Tuck.  ;  Lecanora  frustulosa  (Dicks.)  Mass.,  and  Buellia  pul- 
lata  Tuck.  With  this  much  in  hasty  review  I  shall  pass  to  lo- 
calities more  thoroughly  studied.  However,  I  may  add  here 
better  than  elsewhere  in  my  paper  that  Rinodina  oreina  (Ach.) 
Mass,  and  Lecanora  xanthophana  Nyl.  are  here  and  elsewhere 
in  the  valley  far  more  abundant  than  I  have  ever  found  them  in 
other  regions. 

In  attempting  a  general  comparative  study  of  distribution  in  the 
valley  the  places  that  present  questions  of  greatest  interest  are  the 
vicinities  of  Mankato  and  Granite  Falls,  where  all  sorts  of  sub- 
strata were  examined.  The  two  areas  were  about  equally  well 
studied,  though  the  former,  because  of  the  greater  number  and 


Fink :   LICHENS  OF  THE  MINNESOTA  VALLEY.  279 

less  accessibility  of  rocky  substrata,  required  more  time.  The 
former  area  furnished  151  species  and  varieties  and  the  latter  124. 
A  brief  analysis  of  the  causes  of  the  advantage  in  favor  of  the 
former  region  can  be  best  made  by  a  consideration  of  the  sub- 
joined table,  giving  the  various  substrata  for  both  localities  with 
the  number  of  lichens  most  commonly  found  on  each. 

Numbers  for  Mankato.  For  Grai.ite  Falls. 
Trees                                   60  41 

Rocks  55  54 

Earth  22  17 

Dead  wood  14  12 

A  complete  analysis  introducing  per  cents  as  was  made  in  a 
former  paper  is  not  necessary  since  general  likeness  except  for 
trees  is  apparent  in  the  table.  The  difference  in  richness  then 
is  due  mainly  to  absence  of  large  areas  of  trees  at  Granite 
Falls.  The  slight  differences  in  the  other  three  items  in  the 
table  is  doubtless  due  to  difference  in  moisture,  the  precipitation 
being  30.53  inches  annually  at  Mankato  for  three  years  for 
which  I  could  get  data  and  21.83  inches  annually  at  Granite 
Falls  for  five  years  for  which  data  were  obtained.  Difference 
in  moisture  doubtless  also  accounts  in  small  measure  for  the  ad- 
vantage of  the  Mankato  area  as  to  arboreal  lichens. 

As  to  rocky  substrata  favorable  to  lichen  growth  little  can  be 
definitely  given  by  way  of  comparison.  As  to  kinds  of  rocks 
Mankato  has  an  advantage  in  having  the  sandstone  which  is 
wanting  at  Granite  Falls,  and  also  in  the  great  masses  of  lime- 
stone which  are  replaced  at  Granite  Falls  only  by  the  calcareous 
drift  pebbles  and  a  few  bowlders.  Yet  these  two  advantages  are 
probably  quite  overcome  by  the  great  masses  of  exposed  gran- 
ite at  Granite  Falls,  not  replaced  at  Mankato  in  any  way,  since 
granitic  bowlders  are  equally  abundant  in  both  places. 

Comparing  the  Mankato  vicinity  with  Minneapolis  and  with 
Fayette,  Iowa,  two  areas  compared  in  a  former  portion  of  these 
studies,  we  find  that  it  has  a  much  richer  lichen  flora  than  the 
former  region  which  gave  only  113  lichen  forms  and  probably 
nearly  as  rich  as  the  latter  which  gave  157  lichens  which  one 
could  expect  to  find  in  a  study  of  limited  duration. 

Minnesota  has  now  furnished  more  lichens  than  any  other 
state  in  the  Mississippi  Valley,  having  351  species  and  varieties. 
Illinois  with  249  lichens  being  next  in  order.  Yet  the  fact  that 


280  MINNESOTA    BOTANICAL    STUDIES. 

northeastern  Iowa,  a  portion  of  a  State  not  so  thoroughly  sur- 
veyed and  only  having  226  known  lichens,  has  26  lichens  not 
yet  found  in  Minnesota,  shows  that  the  study  of  Minnesota 
lichens  is  by  no  means  yet  approximately  completed,  since  a 
large  part  of  these  26  rare  or  obscure  lichens  found  already 
within  50  miles  of  the  state  certainly  exist  within  its  bord- 
ers in  the  southeastern  portion,  and  other  unstudied  portions 
of  the  state  may  yet  be  expected  to  bring  additions  to  the 
lichen  flora  in  like  proportion.  A  list  of  these  26  lichens  could 
be  added  with  habitats  to  aid  in  their  discovery  in  southeastern 
Minnesota  but  an  inspection  of  another  paper*  will  give  the 
names  of  them. 

A  study  of  the  table  above,  giving  habitats  and  number  of 
species  for  each,  by  per  cents,  would  give  a  somewhat  larger 
per  cent,  of  lithophytic  lichen  species  for  the  two  areas  consid- 
ered than  a  former  study  exhibited  for  the  Minneapolis  and 
Iowa  localities  and  about  the  same  per  cent,  as  the  lake  Superior 
region.  I  subjoin,  arranged  according  to  habitat,  a  list  of  the 
41  lichens  added  to  the  state  in  this  paper.  From  the  list  it  will 
be  seen  that  more  than  half  of  these  species  are  most  common 
on  rocks,  and  that  the  great  Archean  and  Algonkian  masses 
exposed  throughout  the  upper  valley  alone  produced  one-third 
of  them.  For  convenience  of  reference  to  the  above  statements 
I  shall  now  add  the  table,  placing  rock  species  first,  and  then 
follow  the  list  with  further  discussion. 

New  to  Minnesota  on  Archean  or  Algonkian  rocks. 
Ramalina  polymorpha  (Acn.)  TUCK. 

Parmelia  saxatilis  (L.)  FR.  var.  panniformis  (Acn.)  SCHAER. 
Pyrenopsis  phaeococca  TUCK. 
Pyrenopsis  melambola  TUCK. 
Omphalaria  phyllisca  (WAHL.)  TUCK. 
Leptogium  pulchellum  (AcH.)  NYL. 
Lecanora  sp. 

Lecanora  subfusca  (L.)  ACH.  var.  allophana  ACH. 
Lecanora  cinerea  (PERS.)  NYL.  var.  cinereoalba  var.  nov. 
Rinodina  sophodes  (Acn.)  NYL.  var.  tephraspis  TUCK. 


*  Fink,  B.  Review  of  Lichenological  Studies  in  the  Upper  Mississippi  Valley, 
with  suggestions  for  future  investigations.  In  list  to  be  published  in  Memoirs 
of  the  Torrey  Botanical  Club. 


I'"lllk  :     LICHENS    OF    THE    MINNESOTA    VALLEY.  281 

Rinodina  lecanorina  MASS. 
Urceolaria  actinostoma  PERS. 
Buellia  pullata   TUCK. 

New  to  Minnesota  on  limestone. 
Omphalaria  kansana  TUCK. 
Omphalaria  pulvinata  NYL. 
Collema  plicatile  SCHAER. 
Collema  pustulatum  ACH. 
Lecanora  bookii  (FR.)  TH.  FR. 
Rinodina  bischoffii  (HEPP.)  KOERB. 
Buellia  alboatra  (HOFFM.)  TH.  FR.  var.  saxicola  FR. 
Staurothele  diffractella  (NYL.)  TUCK. 

New  to  Minnesota  on  wood. 
Placodium  ferrugineum  (HUDS.)  HEPP. 
Placodium  ferrugineum  (Huos.)  HEPP.  var.  pollinii  TUCK. 
Cladonia  cristatella  TUCK.  var.  paludicolaTucK. 
Biatora  flexuosa  FR. 
Biatora  suffusaFR. 
Buellia  turgescens  (NYL.)  TUCK. 
Opegrapha  varia  (PERS.)  FR.  var.  pulicaris  FR. 
Arthonia  sp. 

Endocarpon  arboreum  SCHWEIN. 
Pyrenula  gemmata  (Acn.)  NAEG. 
Pyrenula  hyalospora  NYL. 
Pyrenula  quinqueseptata  (NYL.)  TUCK. 
Pyrenula  glabrata  (ACH.)  MASS. 
Pyrenula  megalospora  sp.  nov. 

New  to  Minnesota  on  earth. 
Heppia  despreauxii  (MONT.)  TUCK. 
Heppia  polyspora  TUCK. 
Collema  tenax  (Sw.)  ACH. 
Biatora  decipiens  (EHRH.)  FR. 
Biatora  decipiens  (EHRH.)  FR.  var.  dealbata  AUCT. 

The  list  of  species  new  to  the  state  shows  a  large  number  of 
Pyrenulas,  the  genus  being  unusually  well  represented  in  the 
valley,  especially  at  Mankato.  It  will  also  be  seen  that  the 


282  MINNESOTA    BOTANICAL    STUDIES. 

gelatinous  lichens,  the  Collemei,  are  especially  conspicuous  in 
the  genera  Pyrenops/'s,  Omphalaria,  Collcma  and  Leptogium. 
This  happens  because  part  of  the  valley  is  more  favorable  for 
their  development  as  to  substrata  and  moisture  than  other 
studied  portions  of  the  state.  The  part  of  the  studied  por- 
tion of  the  valley  most  favorable  for  their  development  is  the 
Mankato  vicinity  where  most  of  the  gelatinous  lichens  were 
found.  The  whole  number  of  Collemei  found  in  the  valley  is 
17.  Richness  is  apparent  when  we  add  that  only  four  were 
found  about  Minneapolis,  u  in  the  lake  Superior  region  and 
that  only  16  are  known  in  Iowa. 

It  may  be  added  that  a  large  proportion  of  the  species  added 
to  the  state  flora  are  of  special  interest  for  various  reasons. 
Thus  the  Omphalarias  are  not  commonly  collected  ;  Lecanora 
bookii  (Fr.)  Th.  Fr.  is  a  difficult  lichen  to  detect ;  the  Pyrenulas 
are  difficult  to  distinguish  macroscopically  and  are  therefore 
commonly  overlooked ;  members  of  the  genus  Pyrenopsis  are 
seldom  reported ;  while  Urceolaria  actinostoma  Pers.,  Buellia 
•pullata  Tuck,  and  Heppia  polyspora  Tuck,  are  very  rare  lich- 
ens. Rinodina  lecanorina  Mass,  is  reported  for  the  first  time 
from  North  America,  and  Lecanora  cervina  (Pers.)  Nyl.  var. 
cinereoalba  var.  nov.  is  interesting  because  new. 

It  may  be  noted  in  passing  that  the  region  shows  some  of  the 
Arctic  or  sub- Arctic  species  found  at  Taylor's  Falls  and  already 
discussed  in  a  former  paper.  These  are  Biatora  rufonigra 
Tuck.,  two  forms  of  Buellia  petraa  (Flot.,  Koerb.)  Tuck, 
and  an  Ephebe,  though  not  the  species  reported  from  Taylors 
Falls.  As  in  the  Taylors  Falls  region  the  Buellia  is  the  most 
common  of  these  species  being  a  crustaceous  form  well  adapted 
to  resist  unfavorable  conditions.  The  Biatora  is  next  in  fre- 
quency of  occurrence  and  the  Ephebe,  a  fruticulose  form,  was 
only  seen  once.  So  far  as  I  was  able  to  ascertain  bv  careful 
search  the  foliaceous  forms,  Umbilicaria  and  Nephroma,  found 
at  Taylors  Falls  have  not  succeeded  in  persisting  in  the  Min- 
nesota valley.  This  failure  of  northern  forms  to  persist  so  suc- 
cessfully may  be  accounted  for  perhaps  in  a  very  small  degree 
by  more  southern  position  of  the  area  now  under  consideration, 
but  no  doubt  is  due  much  more  to  climatic  and  edaphic  factors 
which  have  allowed  plant  migrations  to  proceed  northward  more 
rapidly  in  the  Minnesota  valley  than  farther  east  in  the  state 
since  the  last  retreat  of  the  glaciers.  This  matter  has  been 


LICHENS    OF    THE    MINNESOTA    VALLEY.  283 

touched  upon  by  Professor  C.  MacMillan.*  It  is  interesting  to 
note  that  the  strictly  crustaceous  Bucllia  is  the  only  one  of  the 
more  northern  forms  found  in  the  state  which  persists  as  far 
south  as  Pipestone.  Indeed,  its  abundance  here  and  records  of 
occurrence  elsewhere  well  southward  in  low  altitudes  since 
Tuckerman  wrote  lead  to  the  suspicion  that  it  may  not  be  so 
strictly  sub-Arctic  in  distribution  as  I  have  supposed.  It  may 
be  added  that  the  Buellia  is  the  only  one  of  these  northern 
species  persisting  in  the  valley,  which  was  found  on  bowlders  at 
anv  considerable  distance  from  the  large  masses  of  Archean 
and  Algonkian  rocks,  which  are  supposed  to  have  been  exposed 
continuously  since  the  close  of  the  glacial  epoch,  and  that  it 
was  only  found  once  in  very  small  quantity  on  a  bowlder  re- 
mote from  these  larger  masses. 

It  has  been  my  plan  to  introduce  in  each  paper  of  the  series 
some  feature  regarding  distribution  which  could  be  especially 
well  illustrated  by  the  area  under  consideration.  In  the  study 
of  the  Minnesota  valley  and  southwestern  Minnesota  I  was 
able  to  keep  in  mind  a  variety  of  ecologic  factors  and  to  pre- 
serve the  data  necessary  for  their  solution.  This  I  had  pre- 
viously done  in  part  for  several  areas  in  Minnesota  and  Iowa  so 
that  in  the  present  paper  interesting  and  instructive  comparisons 
can  be  made.  Leaving  other  questions,  then,  thus  briefly 
stated,  I  shall  now  pass  to  a  consideration  of  the  lichen  for- 
mations of  the  region,  causes  of  their  peculiar  make-up,  and 
comparisons  with  similar  formations  within  and  outside  the  area 
under  consideration. 

Aside  from  the  purely  scientific  interest  of  the  analysis  to 
follow,  it  has  a  practical  bearing,  in  that  knowledge  of  the  re- 
lation between  ecologic  factors  and  distribution  enables  the  col- 
lector to  predict  in  the  field  about  what  species  of  lichens  he 
may  expect  to  find  in  a  spot  having  a  given  set  of  environmen- 
tal features.  In  the  study  species  rarely  found  in  the  forma- 
tions have  not  been  considered  when  there  appeared  to  be  doubt 
as  to  whether  they  were  collected  on  their  usual  substrata,  and 
rarer  varieties  have  been  omitted  when  showing  the  same  habi- 
tat as  other  forms  of  the  species.  It  will  be  readily  granted 
that  the  commoner  forms  which  give  character  to  the  flora  are 
the  ones  which  should  receive  attention  in  such  a  study.  In 

*  MacMillan,  C.,  •  Observations  on  the  Distribution  of  Plants  Along  the 
Shore  at  Lake  of  the  Woods.  Minn.  Bot.  Stud,  i  :  967.  1897. 


284  MINNESOTA    BOTANICAL    STUDIES. 

the  analysis,  especially  as  to  amount  of  illumination  and  the 
roughness  of  ligneous  substrata,  it  will  be  seen  that  lines  can 
not  be  drawn  very  closely  without  entailing  an  amount  of 
minutiae  which  would  be  confusing  and  therefore  unprofitable. 
With  the  above  brief  statement  as  to  the  main  purpose  of  the 
present  paper,  I  shall  begin  the  consideration  of  lichen  forma- 
tions with  the  most  distinct  ones  with  which  I  am  acquainted, 
viz.,  those  of  the  Sioux  quartzite  at  Pipestone.  These  forma- 
tions are  distinct  because  for  most  part  removed  from  trees  from 
which  lichens  commonly  migrate  to  rocks  nearby,  producing 
tension  lines  and  mixture  of  formations  and  because  the  few 
young  trees  found,  though  large  enough  to  bear  the  foliaceous 
lichens  which  commonly  migrate  to  the  rocks,  have  apparently 
been  isolated  from  larger  areas  of  trees  from  the  beginning  of 
growth  and  scarcely  bear  a  lichen  of  any  kind.  The  rocky 
substratum  is  for  the  most  part  horizontal  and  exposed  to  the 
sun's  rays.  In  a  few  places  occur  perpendicular  rock  expos- 
ures which  are  more  or  less  shaded  by  trees,  overhanging  rocks 
or  north  exposure.  A  few  ombrophytic  lichens  occupy  these 
spots ;  but  they  are  all  strictly  lithophytic  species,  none  of  them 
having,  for  the  reason  stated  above,  migrated  from  trees  as  we 
shall  find  to  be  the  condition  in  a  later  analysis  of  other  similar 
formations.  Below  I  give  first  the  lichen  formation  of  the  hori- 
zontal exposed  rocks  and  second,  that  of  the  more  or  less  shaded 
and  damp  rocks.  Lecanoras  predominate  in  the  formations  on 
exposed  rocks,  which  may  accordingly  be  named  as  follows  : 

Lecanora  formation    of  the  horizontal  exposed  quartzite 

(Pipestone). 

Parmelia  olivacea  (L.)  ACH.  var.  prolixa  ACH.,  C. 
Parmelia  conspersa  (EHRH.)  ACH.,  C. 
Physcia  tribacia  (Acn.)  TUCK.,  C. 
Physcia  caesia  (HOFFM.)  NYL. 
Placodium  elegans  (LINK.)  DC.,  C. 
Placodium  vitellinum  (EHRH.)  NAEG.  and  HEPP. 
Lecanora  rubina  (VILL.)  ACH.,  C. 

Lecanora  rubina  (ViLL.)  ACH.  var.  heteromorpha  ACH.,  C. 
Lecanora  cinerea   (L.)  SOMMERF.,  C. 
Lecanora  xanthophana  NYL.,  C. 


Fink:    LICHENS  OF  THE  MINNESOTA  VALLEY.  285 

Rinodina  oreina  (AcH.)  MASS.,  C. 

Buellia  spuria  (SCHAER.)  ARN.,  C. 

Buellia  pullata  TUCK.,  C. 

Buellia  petraea  (PLOT.,  KOERB.)  TUCK.  var.  montagnaei 
TUCK.,  C. 

Endocarpon  miniatum  (L.)  SCHAER.  var.  complicatum 
SCHAER.,  C. 

The  formation  on  shaded  rocks  may  be  designated  the  Stauro- 
thclc  formation,  after  the  prevailing  genus. 

Staurothele  formation  of  shaded  or  damp  quartzile  (Pipestone). 

Endocarpon  miniatum  (L.)  SCHAER. 
Staurothele  umbrina  (WAHL.)  TUCK.,  C. 
Staurothele  drummondii  TUCK.,  C. 

The  lichen  formations  of  the  pipestone  lying  beside  the  quartz- 
ite  were  studied  to  ascertain  to  what  extent  the  difference  in 
chemical  composition  and  hardness  of  the  rocks  would  influence 
the  distribution  of  lichens,  other  ecologic  factors  being  identical. 
In  the  above  table  I  have  indicated  species  common  to  quartz- 
ite  and  pipestone  by  (C.),  and  the  table  shows  that  only  three 
lichens  were  detected  on  the  quartzite  and  not  on  the  pipestone. 
The  following  three,  all  growing  in  exposed  places,  were  found 
on  the  latter  and  not  on  the  former. 

Placodium  cinnabarrinum  (Acn.)  Auz. 

Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP.  var.  sideritis 
TUCK. 

Lecanora  muralis  (SCHREB.)  SCHAER.  var.  saxicola  SCHAER. 

It  is  worthy  of  note  that  the  differences  are  specific  and  that 
the  formations  are  identical  generically.  The  appearance  of  a 
certain  plant  in  a  particular  set  of  ecological  conditions  is  too 
complicated  a  matter  for  exact  explanation  in  many  instances, 
and  I  can  offer  no  explanation  as  to  why  the  few  plants  occur 
on  one  kind  of  rock  and  not  on  the  other.  Possibly  the  specific 
acid  secreted  by  a  particular  species  acts  more  readily  on  one 
kind  of  rock  than  on  the  other,  but  more  probably  the  cause  is 
other  than  this.  Nor  do  I  suppose  that  I  have  found,  here  or 
in  other  formations  to  be  considered  below,  all  the  lichens  grow- 
ing under  a  particular  set  of  conditions.  Yet  the  common  ones 
which  give  character  to  the  various  formations  were  doubtless 


28G  MINNESOTA    BOTANICAL    STUDIES. 

all  detected  here  as  elsewhere,  and  the  fact  that  15  of  18  were 
found  on  each  kind  of  rock  demonstrates  that  difference  in 
composition  of  rock  in  this  instance  has  produced  little,  if  any 
difference  in  lichen  flora.  A  similar  study  of  lichen  formations 
on  large  rock  areas  of  greater  difference  in  composition  as  granite 
and  limestone  lying  adjacent  would  be  of  special  interest. 

To  complete  the  lichen  formations  of  the  area,  the  earth- 
lichen  formation  must  be  considered.  This  formation  and  simi- 
lar ones  elsewhere  may  be  called  the  Endocarpon  hepalicum  for- 
mations of  exposed  earth  from  a  plant  which  is  found  in  such 
formations  in  all  parts  of  the  state  except  the  lake  Superior 
region. 

Endocarpon  hepaticum  formation  of  exposed  earth  (Pipestone). 
Urceolaria  scruposa  (L.)  NYL. 
Cladonia  pyxidata  (L.)  FR. 
Cladonia  fimbriata  (L.)  FR. 
Cladonia  fimbriata  (L.)  FR.  var.  tubaBformis  FR. 
Biatora  muscorum  (Sw.)  TUCK. 
Endocarpon  hepaticum  ACH. 
Endocarpon  pusillum  HEDW.  var.  garovaglii  KPH. 

The  region  is  a  comparatively  dry  one  because  of  small  pre- 
cipitation of  moisture,  since  the  rocks  lie  high  where  there  is 
little  or  no  standing  water  to  give  moisture  and  because  there 
are  few  trees  to  give  shade.  The  lichen  formations  are  accord- 
ingly rather  poor  in  species,  as  will  appear  in  comparisons  to 
follow  an  analysis  of  similar  formations. 

The  rocky  surfaces  at  Granite  Falls  present  a  much  more 
complex  set  of  conditions  than  those  just  considered,  and  yet, 
for  my  purpose,  they  may  be  classified,  like  the  latter,  into  ex- 
posed surfaces,  usually  horizontal,  and  shaded  surfaces,  usually 
more  or  less  nearly  perpendicular.  I  shall  now  record  these 
formations  in  the  same  order  as  in  the  last  series  ;  but  after  each 
shall  compare  it  with  the  corresponding  formation  at  Pipestone, 
giving,  as  far  as  possible,  the  probable  cause  of  differences. 

Lecanora  formation  of  exposed  (usually  horizontal}  granite 

(Granite  Falls). 

Parmelia  olivacea  (L.)  ACH.  var.  prolixa  ACH. 
Parmelia  conspersa  (EHRH.)  ACH. 


Fink:     LICHENS    OF    THE    MINNESOTA  VAI. I. KY.  287 

Physcia  stellaris  (L.)  TUCK.  var.  apiola  NYL.,  A. 

Physcia  caesia  (HOFFM.)  NYL. 

Placodium  elegans  (LINK.)  DC. 

Placodium  murorum  (HOFFM.)  DC.,  A. 

Placodium  cinnabarrinum  (Aci-i.)  Auz. 

Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP.  var.  sideritis 
TUCK. 

Placodium  vitellinum  (EHRH.)  NAEG.  and  HEPP. 

Lecanora  rubina  (VILL.)  ACH. 

Lecanora  rubina  (VILL.)  ACH.  var.  heteromorpha  ACH. 

Lecanora  muralis  (SCHREB.)  SCHAER.,  A. 

Lecanora  muralis  (SCHREB.)  SCHAER.  var.  saxicola  SCHAER. 

Lecanora  frustulosa  (DICKS.)  MASS.,  A. 

Lecanora  subfusca  (L.)  ACH.  var.  allophana  ACH.,  A. 

Lecanora  subfusca  (L.)  ACH.  var.  coilocarpa  ACH.,  A. 

Lecanora  hageni  ACH.,  A. 

Lecanora  cinerea  (L.)  SOMMERF. 

Lecanora  calcarea  (L.)  SOMMERF.  var.  contorta  FR.,  A. 

Lecanora  xanthophana  NYL. 

Lecanora  cervina  (PERS.)  NYL.  var.  cinereoalba  var.  nov.,  A. 

Lecanora  fuscata  (SCHRAD.)  TH.  FR.,  A. 

Rinodina  oreina  (Acn.)  MASS. 

Rinodina  sophodes  (Acn.)  NYL.,  A. 

Rinodina  lecanorina  MASS.,  A. 

Urceolaria  actinostoma  PERS.,  A. 

Biatora  rufonigra  TUCK.,  A. 

Buellia  spuria  (SCHAER.)  ARN. 

Buellia  pullata  TUCK. 

Buellia  petraea  (FLOT.,  KOERB.)  TUCK. 

Endocarpon  miniatum  (L.)  SCHAER.,  var.  complicatum 
SCHAER. 

Comparing  this  lichen  formation  with  the  similar  ones  of  the 
Sioux  quartzite  and  the  pipestone,  we  find  it  to  contain  all 
lichens  found  on  the  two  except  Physcia  tribacia  (Ach.)  Tuck, 
and  to  contain  fourteen  not  found  on  them,  which  I  have  marked 
as  additions  (A).  The  absence  of  the  one  species  from  the 
Granite  Falls  formation  is  doubtless  an  accident  in  plant  distri- 


288  MINNESOTA    BOTANICAL    STUDIES. 

bution  whose  explanation  would  be  very  difficult  or  impossible  to 
trace ;  but  it  is  quite  remarkable  that  with  this  exception  all  the 
plants  found  in  the  two  formations  sixty  miles  away  should  oc- 
cur in  this  lichen  formation  also,  especially  since  there  could 
have  been  no  rocky  connection  between  the  two  areas  since 
glacial  times.  It  is  not  strange  that  the  exposed  granite  lichen 
formation  at  Granite  Falls  should  be  a  much  richer  one  than  the 
two  exposed  formations  at  Pipestone  combined  ;  for  it  is  a  much 
larger  area,  is  connected  with  a  limestone  lichen  formation  and 
an  epiphytic,  and  a  number  of  swamps  and  ponds  furnish  mois- 
ture along  the  borders.  Indeed  the  presence  of  ten  of  the 
fourteen  additions  may  be  more  or  less  satisfactorily  explained. 
These  I  shall  proceed  to  consider  seriatim. 

Physcia  stellaris  (L.)   TUCK.,    var.    apiola  TUCK. — a  litho- 
phytic  variety  of  a  species  common  on  adjacent  trees. 

Lecanora  frustulosa  (DICKS.)  MASS. — a  northern  lichen   not 
extending  so  far  south  as  Pipestone. 

Lecanora  subfusca  (L.)  ACH.,  var.  alliophana  ACH. — a  variety 
of  a  species  common  on  trees  near  by. 

Lecanora  subfusca  (L.)  ACH.,  var.  coilocarpa  ACH. — as  the  last 
above. 

Lecanora  cervina  (PERS.)  NYL.,  var.  cinereoalba  var.  nov.— 
has  not  been  seen  outside  the  Minnesota  valley. 

Lecanora  calcarea  (L.)  SOMMERF.,  var.  contorta  FR. — a  lichen 
migrating  from  the  limestone  near  by. 

Rinodina  sophodes  (Acn.)  NYL. — found  on  trees  of  the  region 
and  perhaps  migrating  from  them. 

Rinodina  lecanorina  MASS. — a  very  rare  plant  which,  therefore, 
very  probably  does  not  exist  at  Pipestone  or  was  overlooked. 

Urceolaria  actinostoma  PERS. — as  the  last  above. 

BiatorarufonigraTucK. — a  northern  form  not  extending  so  far 
south  as  Pipestone. 

Though  somewhat  confusing  another  similar  lichen  formation 
must  be  introduced  here  for  comparison  as  follows  : 

Lecanora  formation  of  exposed  quartzite  (New  Ulm). 
Parmelia  conspersa  (EHRH.)  ACH.,  CTS. 
Physcia  caesia  (HOFFM.)  NYL.,  CTS. 

Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP.  var.  sideritis 
Tuck.,  CT. 


Fink  :     LICHENS    OF    THE    MINNESOTA    VALLEY.  289 

Placodium  vitellinum  (EiiRH.)  NAEG.  and  HEPP.,  CTS. 
Lecanora  rubina  (VILL.)  ACH.,  CTS. 

Lecanora  rubina  (VILL.)  ACH.  var.  heteromorpha  ACH.,  CS. 
Lecanora  subfusca  (L.)  ACH.,  S. 
Lecanora  varia  (EHRH.)  NYL..  AS. 
Lecanora  cinerea  (L.)  SOMMERF.,  CTS. 
Lecanora  xanthophana  NYL.,  C. 
Rinodina  oreina  (Acn.)  MASS.,  C. 
Rinodina  sophodes  (Acn.)  NYL. 
Biatora  rufonigra  TUCK.,  T. 
Biatora  myriocarpoides  (NYL.)  TUCK.,  A. 
Buellia  spuria  (SCHAER.)  ARN.,  CT. 
Buellia petraea  (FLOT.,  KOERB.)  TUCK.,  CTS. 
Endocarpon     miniatum     (L.)     SCHAER.     var.    complication 
SCHAER.,  C. 

Comparing  the  above  lichen  formation  with  the  similar  ones 
at  Pipestone  and  Granite  Falls  we  find  it  to  contain  only  two 
species  which  are  additions  to  the  three  at  the  two  places  just 
named.  These  I  have  marked  (A).  It  is  about  as  extensive 
an  area  as  the  two  at  Pipestone  combined,  has  about  the  same 
number  of  lichens  as  both  and  has  12  species  (marked  C)  which 
are  common  to  all  the  exposed  rock  lichen  formations  in  the 
area  considered  in  this  paper.  In  general  these  12  species  may 
be  regarded  as  the  most  constant  of  the  exposed  Archean  and 
Algonkian  rock  lichen  formations  of  southwestern  Minnesota. 
As  we.  multiply  areas  of  comparison  and  especially  as  we  introduce 
those  at  a  greater  distance  the  number  of  common  floral  elements 
very  naturally  decreases.  Thus  considering  the  similar  forma- 
tion at  Taylors  Falls,  we  find  only  8  species  (marked  T)  com- 
mon to  it  and  all  the  similar  ones  previously  considered,  and 
passing  to  the  corresponding  formation  at  Gunflint  in  the  lake 
Superior  region,  the  number  found  in  all  these  similar  forma- 
tions in  widely  separated  areas  of  the  state  is  found  to  be  only 
6  (marked  S).  These  6  species  may  be  looked  for  with  con- 
siderable certainty  wherever  such  lichen  formations  are  well  de- 
veloped in  the  state.  Other  elements  will  vary  according  to 
relation  to  other  adjacent  formations,  position  northward  or 
southward  and  in  some  instances  eastward  or  westward  in 
the  state  and  to  various  ecologic  factors  which  cannot  be  enum- 
erated fully. 


290  MINNESOTA    BOTANICAL    STUDIES. 

We  may  now  turn  to  the  lichen  formation  of  shaded  or  damp 
rocks  at  Granite  Falls.  This  includes  some  flat  rock  surfaces 
somewhat  shaded  or  simply  wet  part  of  the  time,  as  well  as  the 
perpendicular  shaded  surfaces.  I  shall  divide  the  formation 
into  three  parts — species  naturally  belonging  to  the  rocks,  those 
which  have  probably  migrated  from  the  trees  near  at  hand  and 
those  which  have  probably  migrated  from  the  earth.  Here  and 
in  another  formation  we  have  a  mixture  of  elements,  hence  the 
following  name  is  proposed  : 

Mixed  formation  of  shaded  (or  damp]  granite  (Granite  Falls). 

A.  PROBABLY  NATURALLY  BELONGING  TO  THE  ROCKS. 
Ramalina  polymorpha  (Acn.)  TUCK. 

Ramalina  calicaris  (L.)  FR.  var.  farinacea  SCIIAER. 

Pannaria  microphylla  (Sw.)  DELIS. 

Pannaria  languinosa  (AcH.)  KOERB. 

Omphalaria  phyllisca  (WAHL.)  TUCK. 

Collema  furvum  (Acn.)  NYL. 

Leptogium  lacerum  (Sw.)  FR. 

Endocarpon  muriatum  (L.)  SCHAER. 

Staurothele  umbrina  (WAHL.)  TUCK. 

Staurothele  diffractella  (NYL.)  TUCK. 

Staurothele  drummondii  TUCK. 

B.  NEAR  TREES  AND  PROBABLY  MIGRATED  FROM  THEM. 
Parmelia  cetrata  ACH. 

Parmelia  crinita  ACH. 

Parmelia  borreri  TURN. 

Parmelia  borreri  TURN.  var.  hypomela  TUCK. 

Parmelia  saxatilis  (L.)  FR. 

Parmelia  saxatilis  (L.)  FR.  var.  sulcata  NYL. 

Parmelia  saxatilis  (L.)  FR.  var.  panniformis  (Acn.)  SCHAEK. 

Parmelia  caperata  (L.)  ACH. 

Physcia  speciosa  (WULF.,  ACH.)  NYL. 

Physcia  pulverulenta  (SCHREB.)  NYL. 

Physcia  obscura  (EHRH.)  NYL. 

Pyxine  sorediata  FR. 

Leptogium  myochroum  (EHRH.,  SCIIAER.)  TUCK. 


Fink  :    LICHENS  OF  THE  MINNESOTA  VAM.KY.  291 

Placodium  aurantiacum  (LiGirrr.)  NAEG  and  HEPP. 
Biatora  fuscorubella  (HOFFM.)  TUCK. 

C.     SPECIES  WHICH  HAVE  PROBABLY  MIGRATED  FROM  EARTH. 
Peltigera  rufescens  (NECK.)  HOFFM. 
Peltigera  canina  (L.)  HOFFM. 

Of  the  three  parts  of  the  formation  under  consideration  only 
the  first  can  be  compared  with  the  similar  formation  at  Pipe- 
stone,  and  we  find  besides  the  3  species  of  the  Pipestone  for- 
mation, 8  additional  forms  as  a  result  of  greater  areas  studied, 
more  moist  conditions  near  the  Minnesota  river,  and  where 
abundant  ponds  and  marshes  situated  near  the  rocks  give  moist- 
ure, and  where  trees  are  numerous  in  some  parts  of  the  area 
and  increase  the  shade.  I  must  add  that  presence  of  the 
Ramalinas  here,  and  their  absence  from  shaded  rocks  at  Pipe- 
stone  leads  to  the  suspicion  that  they  may  have  sprung  from 
RamaUna  calicaris  (L.)  Fr.  of  the  region,  migrating  from  trees 
to  rocks  and  acquiring  the  varietal,  and  in  one  instance  the 
specific  characters  as  an  adaptation  to  changed  environment. 
The  question  is  as  to  whether  these  lichens  are  sufficiently  plastic 
to  acquire  such  new  characters  since  trees  have  grown  in  the 
valley  in  post-glacial  time.  I  can  only  say  that  I  believe  that 
they  may  be,  and  that  it  is  quite  as  likely  that  the  two  Rama- 
linas should  be  placed  in  the  second  division  of  the  formation 
as  in  the  first. 

As  to  plants  of  the  second  portion  of  the  formation,  which  I 
have  designated  as  having  probably  migrated  from  trees,  in 
some  instances  they  are  locally  more  abundant  and  luxuriant  on 
the  rocks  than  on  trees.  Hence  a  hasty  consideration  would 
lead  to  the  conclusion  that  they  have  not  migrated.  But  the 
luxuriant  condition  obtains  on  the  rocks  in  Parmelia  borreri 
Turn.,  a  lichen  seldom  seen  on  rocks  elsewhere,  and  many  of 
these  lichens  grow  on  mossy  rocks  where  lichens  are  commonly 
large.  Also  it  is  to  be  taken  into  account  that  these  lichens  are 
those  usually  found  on  large  trees  with  rough  bark.  The  larger 
trees  were  for  most  part  destroyed  years  ago  by  man  or  fires, 
and  these  lichens,  formerly  common  on  trees,  are  preserved  on 
rocks  better  than  on  the  less  permanent  trees.  Hence  some  of 
them  are  more  common  now  on  the  rocks  than  on  the  trees, 
which  are  for  most  part  second  growth  and  not  large.  The 


292  MINNESOTA    BOTANICAL    STUDIES. 

third  division,  consisting  of  two  Peltigeras^  scarcely  needs  any 
special  consideration. 

I  shall  next  consider  the  similar  shaded  rock  formation  at 
New  Ulm,  which  may  be  divided  into  those  lichens  naturally 
belonging  to  the  rocks  and  those  probably  migrating  from  trees. 

Mixed  lichen  formation  of  shaded  rocks  (New  Ulm). 

A.  NATURALLY  BELONGING  TO  THE  ROCKS. 
Pannaria  languinosa  (AcH.)  KOERB. 
Collema  flaccidum  ACH. 

Collema  furvum  (Acn.)  NYL. 

B.  NEAR  TREES  AND  PROBABLY  MIGRATED  FROM  THEM. 
Theloschlstes  lychneus  (NYL.)  TUCK.,  CTS. 
Parmelia  crinita  ACH.,  CTS. 

Parmelia  borreri  TURN.,  CTS. 
Parmelia  saxatilis  (L.)  FR.,  CTS. 

Parmelia  saxatilis  (L.)  FR.  var.  panniformis  (Acn.)  SCHAER., 
C. 

Parmelia  caperata  (L.)  ACH.,  CTS. 
Physcia  speciosa  (WULF.  ACH.)  NYL.,  CTS. 
Physcia  pulverulenta(ScHREB.)  NYL.,  CTS. 
Physcia  stellaris  (L.)  TUCK.,  TS. 
Physcia  obscura  (EHRH.)  NYL.,  CTS. 

As  to  the  shaded  rock  lichen  formations  of  the  region  sur- 
veyed considering  only  plants  naturally  belonging  to  the  rocks, 
there  is  not  a  single  lichen  that  is  common  to  all  of  them.  Pan- 
naria languinosa  (Ach.)  Koerb.  is  the  most  constant  element 
of  such  formations,  which  as  a  whole  might  be  named  for  this 
plant  were  it  not  quite  as  common  in  shaded  limestone  forma- 
tions otherwise  quite  different  from  any  of  those  on  the  rocks 
under  consideration  at  present.  Of  the  lichens  of  the  shaded 
rock  formation  at  New  Ulm,  which  have  probably  migrated 
from  trees,  the  nine  marked  common  (C),  may  be  taken  as  the 
ones  most  commonly  occurring,  as  they  were  found  also  at 
Granite  Falls  in  the  similar  formation.  Those  marked  (T)  all 
but  one  of  the  nine,  occur  in  the  similar  formation  at  Taylors 
Falls.  Other  elements  vary  more  with  change  in  various  eco- 
logic  factors.  The  similar  partial  formation  was  noted  at 
Grand  Portage,  especially  on  the  island,  and  adding  those 


Fink  :     LICHENS    OF    THE    MINNESOTA    VALLEY.  293 

lichens  (S)  of  it  found  in  the  corresponding  ones  considered 
above,  subtracts  none  from  the  number  of  common  species. 
Therefore,  these  eight  lichens  may  be  regarded  as  the  elements 
of  that  portion  of  the  shaded  rock  lichen  formations  which  have 
probably  migrated  from  trees,  most  widely  occurring  in  such 
formations  over  the  state.  Only  one  day  was  spent  in  study- 
ing the  New  Ulm  formations.  A  second  day  would  have  added 
somewhat  to  the  list,  yet  doubtless  all  the  dominant  lichen  floral 
elements  were  secured. 

Without  entering  into  a  detailed  analysis,  it  will  appear  from 
an  inspection  of  the  lichens  composing  the  formations  for  shaded 
and  for  exposed  rocks  that  the  species  occurring  in  the  former 
are  for  most  part  foliaceous  or  fruticulose  types,  while  those 
given  for  the  latter  are  in  general  crustaceous,  or  if  foliaceous, 
at  least  closely  prostrate  on  the  rocks.  This  is  what  would  be 
expected,  since  shade  favors  better  development  of  thallus,  so 
that  those  species  showing  good  thalli  crowd  out  the  other 
species  in  shaded  places,  or  when  unshaded  become  shaded  with 
the  growth  of  trees. 

Next  in  order  come  the  earth  lichen  formations  of  the  rocky 
areas  of  Granite  Falls  and  New  Ulm.  I  shall  first  record  the 
exposed  formations  for  the  two  localities  and  compare  with  the 
similar  formation  already  recorded  for  Pipestone.  Then  will 
follow  the  lichen  formations  of  shaded  earth  at  the  first  two  sta- 
tions, which  is  scarcely  developed  at  Pipestone.  A  consid- 
eration of  calcareous-earth  lichen  formations  follows,  the  pres- 
ent being  formations  of  non-calcareous  earth. 

Endocarpon     hepaticum    lichen    formation    of  exposed   earth 

(Granite  Falls). 

Heppia  despreauxii  (MONT.)  TUCK. 
Urceolaria  scruposa  (L.)  NYL. 
Cladonia  pyxidata  (L.)  FR. 
Biatora  muscorum  (Sw.)  TUCK. 
Biatora  icterica  MONT. 
Endocarpon  hepaticum  ACH. 
Endocarpon  pusillum  HEDW.  var.  garovaglii  KPH. 

Endocarpon    hepaticum    lichen   formation    of   exposed   earth 

(New  Ulm). 

Cladonia  pyxidata  (L.)  FR.,  CTS. 


294  MINNESOTA    BOTANICAL    STUDIES. 

Cladonia  turgida  (EHRH.)  HOFFM. 

Biatora  uliginosa  (SCHRAD.)  FR. 

Endocarpon  hepaticum  ACH.,  CT. 

Endocarpon  pusillum  HEDW.  var.  garovaglii  KPH.,  C. 

Comparing  these  lists  with  the  one  given  for  the  correspond- 
ing formation  at  Pipestone,  we  find  three  common  lichens  which 
are  marked  (C)  in  the  list  above.  Two  of  these  marked  (T)  are 
also  found  in  the  similar  formation  at  Taylors  Falls,  and  one 
marked  (S)  is  common  in  like  formations  in  the  lake  Superior 
region.  This  plant  is  the  most  constant  element  in  the  exposed 
earth  lichen  formations  of  the  State,  and  I  should  be  disposed  to 
name  these  Cladonia  pyxidata  lichen  formations,  were  it  not 
that  the  plant,  though  commonly  present  in  exposed  stations, 
thrives  better  in  shaded  ones.  I  must  here  emphasize  that  these, 
as  well  as  the  calcareous-earth  lichen  formations,  grow  on  earth 
in  rocky  places  where  larger  vegetation  is  scanty  and  scattered. 

Next  in  order  come  lichen  formations  of  shaded  earth,  partly 
composed  of  plants  which  grow  also,  though  not  so  well,  in  un- 
shaded places.  From  their  dominant  elements,  these  may  be 
designated  as  follows : 

Cladonia- Peltiger a   lichen  formation    of  shaded   earth 
(Mankato). 

Peltigera  rufescens  (NECK.)  HOFFM. 

Peltigera  canina  (L.)  HOFFM. 

Peltigera  canina  (L.)  HOFFM.  var.  sorediata  SCHAER. 

Collema  pulposum  (BERNH.)  NYL. 

Collema  tenax  (Sw.)  ACH. 

Cladonia  pyxidata  (L.)  FR. 

Cladonia  fimbriata  (L.)  FR. 

Cladonia  gracilis  (L.)  FR. 

Cladonia  gracilis  (L.)  FR.  var.  verticillata  FR. 

Cladonia-Peltigera  lichen  formation  of  shaded  earth  (Granite 

'  Falls). 

Peltigera  rufescens  (NECK.)  HOFFM.,  CT. 

Peltigera  canina  (L.)  HOFFM.,  CTS. 

Peltigera  canina  (L.)  HOFFM.  var.  sorediata  SCHAER.,  CTS. 

Collema  pulposum  (BERNH.)  NYL.,  CT. 


Fink :    LICHENS  OF  THE  MINNESOTA  VALLEY.  '_M>5 

Cladonia  pyxidata  (L.)  FR.,  CTS. 

Cladonia  fimbriata  (L.)  FR.,  C. 

Cladonia  fimbriata  (L.)  FR.  var.  tubasforrais  FR.,  TS. 

Cladonia  gracilis  (L.)  NYL.,  CTS. 

Cladonia  gracilis  (L.)  NYL.  var.  verticillata  FR.,  CTS. 

These  two  formations  are  remarkably  similar,  having  8  com- 
mon forms  (C)  of  a  total  of  nine  lichens  in  each  formation. 
Including  the  similar  formation  at  Taylors  Falls  (T)  we  still 
have  7  lichens  common  to  the  similar  formations  for  a  large 
part  of  Minnesota,  and  extending  the  observation  to  the  similar 
formation  on  Grand  Portage  island  in  the  lake  Superior  (S) 
region,  we  yet  have  6  lichens  common  to  such  formations  selected 
from  widely  separated  areas  in  the  State.  This  is  the  first  kind 
of  formation  thus  far  considered  which  is  found  in  the  Minne- 
apolis area  studied.  Therefore  data  from  this  region  have  not 
been  introduced  thus  far.  Their  use  in  the  present  considera- 
tion would  not  decrease  the  number  of  common  elements,  and 
I  shall  not  add  them.  The  three  rarer  Cladonias  of  the  region 
under  consideration  in  the  present  paper,  Cladonia  symphycarpia 
Fr.,  Cladonia  mitrula  Tuck,  and  Cladonia  cariosa  (Ach.) 
Spreng.  have  been  purposely  omitted,  as  there  is  yet  doubt  as 
to  whether  their  adaptation  is  ombrophytic. 

As  to  the  nature  of  the  lichens  composing  these  earth  lichen 
formations,  it  is  apparent  that  those  of  the  shaded  earth  forma- 
tions are  as  a  whole  more  foliaceous  or  fruticulose  and  better 
developed  as  to  thallus  than  those  of  the  exposed  earth  forma- 
tion. The  explanation  is  of  course  the  same  as  that  already 
given  for  exposed  and  shaded  rock  lichen  formations. 

I  shall  now  consider  the  one  remaining  earth  lichen  formation 
at  Granite  Falls  and  compare  it  with  a  similar  one  in  another 
region.  It  is  that  of  the  earth  among  the  calcareous  drift  peb- 
bles and  small  boulders  on  hill  sides.  From  the  calcareous 
nature  of  the  earth  and  the  presence  of  a  Biatora  seldom  seen 
elsewhere  than  in  such  formations,  the  following  name  has  sug- 
gested itself. 

Biatora  decipiens  lichen  formation  of  exposed  calcareous  earth 

(Granite  Falls). 

Heppia  despreauxii  (MONT.)  TUCK. 
Heppia  polyspora  TUCK. 


296  MINNESOTA    BOTANICAL    STUDIES. 

Urceolaria  scruposa  (L.)  NYL. 

Biatora  muscorum  (Sw.)TucK. 

Biatora  decipiens  (EHRH.)  FR. 

Biatora  decipiens  (EHRH.)  FR.  var.  dealbata  AUCT. 

Endocarpon  hepaticum  ACH. 

Some  of  the  plants  of  this  formation  have  been  found  at 
Mankato  and  also  at  Minneapolis,  but  the  formation  is  not  well 
developed  at  either  place.  However,  it  is  beautifully  developed 
at  Fayette,  Iowa,  and  because  of  its  remarkable  similarity  there 
to  the  Granite  Falls  formation  about  two  hundred  miles  distant, 
I  give  it  below  for  the  sake  of  comparative  study. 

Biatora  decipiens  lichen  formation  of  exposed  calcareous  earth 

(Fayette,  Iowa). 

Heppia  despreauxii  (MONT.)  TUCK.,  C. 

Urceolaria  scruposa  (L.)  NYL.,  C. 

Biatora  muscorum  (Sw.)  TUCK.,  C. 

Biatora  decipiens  (EHRH.)  FR.,  C. 

Biatora  decipiens  (EHRH.)  FR.  var.  dealbata  AUCT.,  C. 

Biatora  fossarum  (DuF.)  MONT. 

Endocarpon  hepaticum  ACH.,  C. 

It  will  be  seen  that  the  two  formations  are  identical  except 
that  each  one  contains  one  species  not  found  in  the  other. 
Again,  this  slight  difference  becomes  less  significant  when  it  is 
stated  that  each  of  these  two  plants  not  found  in  both  formations 
is  rather  rare  in  the  formation  in  which  it  occurs.  The  six 
lichens  common  to  both  formations  I  have  indicated  in  the  Fay- 
ette list  (C).  In  both  localities  the  formations  are  formed  on 
hill  sides  and  seen  to  be  somewhat  better  developed  on  south- 
ward than  on  northward  slopes.  I  have  not  seen  similar  forma- 
tions well  developed  elsewhere,  but  it  is  probable  that  they 
reach  their  best  development  on  unshaded  hill  sides  where  other 
vegetation  is  scanty  and  where  the  lichens  are  washed  with 
lime-impregnated  water  flowing  down  the  slope  during  rains. 
Biatora  decipiens  (Ehrh.)  Fr.  and  Endocarpon  hepaticum  Ach. 
are  the  most  common  plants  of  these  formations,  but  the  latter 
is  quite  as  common  in  another  formation  of  non-calcareous  earth, 
which  I  have  named  for  it,  not  confined  to  hill  sides. 


Fink:     LICHENS    OF    THE    MINNESOTA    VALLEY.  297 

Closely  related  to  the  above  formations  are  two  occupying 
the  same  areas  and  named  for  a  lichen  almost  wholly  confined 
to  them.  They  follow  below  : 

Lecanora  calcarca  contorta  lichen  formation  of  exposed  lime- 
stone pebbles  (Granite  Falls). 

Placodium  vitellinum  (EHRH.)  NAEG.  and  HEPP.  var.  aurel- 
lum  ACH. 

Lecanora  calcarea  (L.)  SOMMERF.  var.  contorta  FR. 

Lecanora  privigna  (Acn.)  NYL. 

Lecanora  privigna  (Acn.)  NYL.  var.  pruinosa  AUCT. 

Endocarpon  pusillum  HEDW. 

Verrucaria  muralis  ACH. 

Staurothele  diffractella  (NYL.)  TUCK. 

Like  the  last,  this  formation  is  not  well  developed  in  other 
studied  portions  of  Minnesota,  and  I  shall  give  the  similar  one 
for  Fayette,  Iowa,  for  comparison. 

Lecanora  calcarea  contorta  lichen  formation  of  exposed  lime- 
stone pebbles  (Fayette,  Iowa). 

Placodium  cinnabarinum  (AcH.)  Auz. 

Placodium  vitellinum  (EHRH.)  NAEG.  &  HEPP.  var.  aurellum 
ACH.,  C. 

Lecanora  muralis  (SCHREV.)  SCHAER.  var.  versicolor  FR. 

Lecanora   calcarea  (L.)  SOMMERF. 

Lecanora  calcarea  (L.)  SOMMERF.  var.  contorta  Fr.,  C. 

Lecanora  privigna  (Acn.)  NYL.,  C. 

Rinodina  bischoffii  (HEPP.)KOERB. 

Biatora  russellii  TUCK. 

Endocarpon  pusillum  HEDW.,  C. 

Verrucaria  nigrescens  PERS. 

Verrucaria  muralis  ACH.,  C. 

Lichens  common  to  the  two  formations  are  marked  (C)  in  the 
Fayette  list,  and  comparison  shows  marked  similarity  in  the  two 
formations  about  200  miles  distant,  except  that  'the  latter  is  con- 
siderably better  developed  than  the  former.  This  is  as  would 
be  expected  when  we  consider  that  the  Iowa  region  is  one  where 
limestones  abound,  while  the  Minnesota  is  one  in  which  the 
limestone  pebbles  are  those  transported  in  glacial  drift  and  are 


298  MINNESOTA    BOTANICAL    STUDIES. 

less  numerous.  All  the  species  of  these  formations,  except  the 
Biatoras,  .have  been  found  elsewhere  in  Minnesota,  but  not  ag- 
gregated into  definite  formations. 

Comparing  the  last  two  series  of  formations,  viz.,  those  of 
calcareous  earth  and  those  of  drift  pebbles  of  the  same  areas, 
it  will  be  noted  that  the  former,  because  of  their  position  on 
dry  hill-sides,  consist  as  a  whole  of  lichens  having  small  foli- 
aceous  or  granular  thalli,  while  those  on  the  yet  dryer  and 
harder  calcareous  pebbles  are  almost  entirely  made  up  of  strictly 
crustaceous  plants. 

The  formations  of  exposed  and  shaded  limestone  bluffs  come 
next  in  natural  order,  and  the  analysis  is  difficult,  since  some  of 
the  lichens  of  these  formations  grow  about  equally  well  in  sun- 
shine and  shade.  These  1  shall  indicate  by  an  interrogation 
point  (?).  From  the  prevalence  of  gelatinous  lichens  they  may 
be  named  as  follows  : 

Gelatinous  lichen  formation  of  shaded  (or  damp)  limestone 
bluffs  (Mankato). 

Pannaria  nigra  (HUDS.)  NYL. 

Pannaria  languinosa  (Acn.)  KOERB. 

Omphalaria  kansana  TUCK.  ? 

Omphalaiia  pulvinata  NYL.  ? 

Collema  plicatile  SCHAER. 

Collema  pustulatum  ACH. 

Leptogium  lacerum  (Sw.)  FR. 

Placodium  citrinum  (HOFFM.)  LEIGHT. 

Biatora  inundata  FR. 

Buellia  alboatra  (HOFFM.)  TH.  FR.  var.  saxicola  FR. 

Endocarpon  miniatum  (L.)  SCHAER. 

Staurothele  umbrina  (\VAHL.)  TUCK. 

Similar  formations  do  not  exist  in  other  surveyed  portions  of 
Minnesota,  except  at  Minneapolis,  where  the  development  is 
poor.  It  is  as  follows  : 

Gelatinous  lichen  formation  of  shaded  (or  damp]   calcareous 
rocks  (Minneapolis). 

Pannaria  nigra  (HUDS.)  NYL.,  C. 
Pannaria  languinosa  (Acn.)  KOERB.,  C. 


Fink:    LICHENS  OF  THE  MINNESOTA  VALLEY.  290 

Omphalaria  sp. 

Leptogium  lacerum  (S\v.)  FR.,  C. 
Endocarpon  miniatum  (L.)  SCHAER.,  C. 
Placodium  citrinum  (HOFFM.)  LEIGHT.,  C. 

The  plants  of  the  Minneapolis  list  are  all  but  one  common  (C) 
to  both  formations  and  may  be  regarded  as  characteristic  of 
such  formations.  Since  the  last  formation  is  poorly  developed, 
I  may  add  the  similar  one  for  Fayette,  Iowa,  which  is  better 
developed  than  either  of  the  above. 

Gelatinous   lichen   formation    of  shaded   (or  damp]   calcareous 

rocks   (Fayette,   Iowa). 
Pannaria  nigra  (Huos.)  NYL. 
Pannaria  languinosa  (Acn.)  KOERB. 
Omphalaria  pulvinata  NYL.  ? 
Omphalaria  umbella  TUCK.  ? 
Omphalaria  sp. 
Collema  plicatile  SCHAER.  ? 
Collema  furvum  (Acn.)  NYL.  ? 
Collema  pustulatum  ACH.? 
Leptogium  lacerum  (Sw.)  FR. 
Leptogium  chlorometum  (S\v.)  NYL. 
Placodium  citrinum  (HOFFM.)  LEIGHT. 
Biatora  trachona  FLOT. 

Buellia   alboatra  (HOFFM.)  TH.  FR.  var.  saxtcola  FR. 
Endocarpon  miniatum  (L.)  SCHAER.  ? 
Staurothele  umbrina  (WAHL.)  TUCK.? 

The  introduction  of  the  Fayette  formation  is  of  special  interest 
for  the  following  reason.  The  first  Minnesota  formation  is  a 
mile  back  from  the  Minnesota  river  on  a  bluff  along  which  the 
river  once  flowed,  but  which  now  is  left  dry  except  for  the  trees 
which  overhang  it  and  shade  the  lichens  of  the  formation. 
The  Fayette  formation  is  on  a  bluff  at  the  water's  edge,  and  the 
plants  are  growing  within  one  to  ten  feet  of  the  water.  Doubt- 
less this  in  part  causes  the  greater  richness.  The  Mankato  for- 
mation is  an  interrupted  one,  none  of  the  plants  persisting  in 
wholly  unshaded  spots.  The  .Fayette  formation  on  the  other 
hand,  extends  for  miles,  without  complete  interruption,  wher- 


300  MINNESOTA    BOTANICAL    STUDIES. 

ever  the  bluffs  exist.  With  the  greater  amount  of  moisture  at 
the  water's  edge,  some  of  the  plants  of  the  Fayette  formation 
grow  well  in  sunshine  and  even  on  south  exposures.  These  I 
have  indicated  by  an  interrogation  point  (?).  These,  for  most 
part  gelatinous  lichens,  require  a  good  amount  of  moisture  ;  and 
if  growing  far  from  water  seek  shade  for  it.  In  the  Fayette 
locality  many  trees  have  been  cut  recently  along  the  bluffs  so 
that  the  plants  are  more  exposed  than  formerly.  The  Minne- 
apolis list  can  be  considered  a  formation  only  in  the  sense  of  a 
group  of  plants  growing  under  like  conditions,  for  owing  to 
somewhat  dryer  climate  the  formation  is  poorly  developed  as  to 
individuals  and  may  be  designated  as  a  scattered  formation, 
only  one  or  two  of  the  species  usually  growing  in  one  limited 
area,  along  the  bluffs  and  long  stretches  of  bluff  between  these 
areas  frequently  not  bearing  a  single  plant  of  the  formation. 

Next  in  order  comes  the  lichen  formation  of  exposed  lime- 
stone bluffs,  which  I  shall  designate  as  follows  from  the  presence 
of  a  large  proportion  of  angiocarpous  lichens. 

Angiocarpous  lichen  formation  of  limestone  bluffs  (Mankato). 

Theloschistes  lychneus  (NYL.)  TUCK. 
Placodium  elegans  (LINK.)  DC. 

Placodium  vitellinum  (EHRH.)  NAEG.  and  HEPP  var.  aurel- 
lum  ACH. 

Placodium  aurantiacum  (LIGHT.)  NAEG.  and  HEPP. 
Lecanora  hageni  ACH. 
Lecanora  erysibe  NYL.  ? 
Endocarpon  pusillum  HEDW.  ? 
Endocarpon  miniatum  (L.)  SCHAER.  ? 
Staurothele  diffractella  (NYL.)  TUCK. 
Verrucaria  fuscella  FR. 
Verrucaria  nigrescens  PERS. 
Verrucaria  muralis  ACH. 

I  might  add  similar  formations  from  Minneapolis  and  Fayette, 
Iowa ;  but  the  analysis  is  very  uncertain  so  that  the  comparisons 
could  have  little  value. 

I  shall  now  consider  the  sandstone  bluff  formations  of  certain 
localities,  simply  designating  them  as  formations  of  damp  sand- 
stone since  they  are  found  along  streams  where  the  rocks  are 


Fink  :     LICHENS    OF    THE    MINNESOTA    VALLEY.  301 

well  supplied  with  moisture.  The  first  of  the  formations  is 
almost  completely  shaded,  but  the  second  is  only  partially 
shaded,  moisture,  the  thing  really  sought  by  the  plants,  being 
sufficient  in  more  or  less  exposed  spots  so  that  the  less  ombro- 
phytic  plants  of  the  group  thrive  twenty  or  thirty  feet  from 
the  water's  surface,  and  even  the  more  shade-loving  ones  are 
found  in  exposed  spots  nearer  the  water.  I  shall  now  record 
the  formations  as  follows,  designating  the  less  ombrophytic 
plants  of  the  second  formation  thus  (  ?).  For  these  formations 
I  suggest  the  following  name  from  a  plant  almost  wholly  con- 
fined to  them  in  Minnesota. 

Usnca  barbata  rubiginea  lichen  formation  of  damp  sandstone 
bluffs  (Minneopa  Falls). 

Ramalina  calicaris  (L.)  FR.  var.  farinacea  SCHAER. 

Usnea  barbata  (L.)  FR.  var.  hirta  FR. 

Usnea  barbata  (L.)  FR.  var.  rubiginea  MICHX. 

Peltigera  canina  (L.)  HOFFM.  var.  sorediata  ACH. 

Leptogium  chloromelum  (Sw.)  NYL. 

Pannaria  languinosa  (Acn.)  KOERB. 

Cladonia  furcata  (HUDS.)  FR. 

Cladonia  furcata  (Huos.)  FR.  var.  racemosa  FR. 

Urceolaria  scruposa  (L.)  NYL. 

Usnea  barbata  rubiginea  lichen  formation  of  damp  sandstone 
bluffs  (Minneapolis). 

Ramalina  calicaris  (L.)  FR.  var.  farinacea  SCHAER.,  C. 

Us nea  barbata  (L. )  FR.  var.  hirta  FR.,  C. 

Usnea  barbata  (L.)  FR.  var.  rubiginea  MICHX.?  C. 

Parmelia  conspersa  (EHRH.)  ACH.  ?  T. 

Peltigera  canina  (L.)  HOFFM.  var.  sorediata  SCHAER.,  CE.. 

Pannaria  languinosa  (Acn.)  KOERB.,  C. 

Lecanora  subfusca  (L.)  Acn.  var.  coilocarpa  ACH.  ?  T. 

Urceolaria  scruposa  (L.)  NYL.?  C. 

Cladonia  caespiticia  (PERS.)  FL.,  T. 

Cladonia  cornucopioides  (L.)  FR.?  E. 

Endocarpon  pusillum  (HEDW.)  var.  garovaglii  KPH.,  E. 


302  MINNESOTA    BOTANICAL    STUDIES. 

Comparing  the  two  formations  we  find  six  common  lichens  of 
a  total  of  nine  recorded  for  the  first  and  eleven  for  the  second. 
Similar  formations  occur  at  Pictured  Rocks,  Iowa,  and  at  Rap- 
idan,  but  I  shall  not  multiply  lists.  As  in  the  instance  of  cer- 
tain formations  on  shaded  granite  or  quartzite  recorded  above, 
both  of  these  formations  are  more  or  less  mixed,  being  made 
up  of  lichens  strictly  lithophytic  in  adaptation  and  of  others 
which  have  doubtless  wandered  from  trees  or  from  earth.  As 
I  have  not  been  able  to  study  such  sandstone  bluffs  at  a  distance 
from  trees,  I  have  not  attempted  a  definite  analysis  of  these 
more  limited  formations  as  I  did  for  the  formations  of  the  shaded 
granite  and  quartzite,  but  have  simply  indicated  in  the  second 
list  those  which  have  probably  wandered  from  trees  by  (T)  and 
those  from  earth  by  (E).  I  have  omitted  from  these  sandstone 
formations  some  of  the  rarer  plants  which  I  should  have  in- 
cluded had  I  attempted  an  analysis  of  these  mixed  formations. 

I  shall  now  proceed  to  the  two  formations  of  trees,  viz.,  that 
of  rough  barked  trees  and  that  of  trees  having  smooth  bark. 
The  distinctions  are  difficult  in  some  instances  as  certain  species 
grow  in  both  habitats.  Consequently,  as  in  some  instances,  in 
formations  previously  considered,  some  plants  are  recorded  for 
more  than  one  formation.  Moreover,  it  must  be  added  that 
some  of  those  recorded  for  rough  barked  trees  frequently  seek 
the  smoother  portions  of  the  bark.  The  subfamily  Parmeleiia 
especially  well  developed  in  the  rough  bark  formations,  which 
may  accordingly  be  named  as  follows  : 

Parmelei  lichen  formation  of  trees  -with  rough  bark  (Mankato). 
Ramalina  calicaris  (L.)  FR.  var.  fraxinea  FR.,  G. 
Ramalina  calicaris  (L.)  FR.  var.  fastigiata  FR.,  G. 
Theloschistes  chrysopthalmus  (L.)  NORM.,  G. 
Theloschistes  polycarpus  (EHRH.)  TUCK.,  G. 
Theloschistes  lychneus  (NYL.)  TUCK.,  G. 
Theloschistes  concolor  (DICK.)  TUCK.,  G. 
Parmelia  perforata  (JACK.)  ACH. 
Parmelia  crinita  ACH.,  G. 
Parmelia  borreri  TURN.,  G. 
Parmelia  tiliacea  (HOFFM.)  FLOERK.,  G. 
Parmelia  saxatilis  (L.)  FR. 
Parmelia  caperata  (L.)  ACH.,  G. 


Fink  :     LICHENS    OF    THE    MINNESOTA    VALLEY.  303 

Physcia  granulifera  (Acn.)  TUCK.,  G. 

Physcia  pulverulenta  (SCHREB.)  NYL.,  G. 

Physcia  stellaris  (L.)  TUCK.,  G. 

Physcia  tribacia  (Acn.)  TUCK. 

Physcia  obscura  (EHRI-I.)  NYL.,  G. 

Physcia  adglutinata  (FLOERK.)  NYL. 

Collema  pycnocarpum  NYL.,  G. 

Collema  flaccidum  ACII. 

Leptogium  myochroum  (EHRH.,  SCHAER.)  TUCK. 

Placodium  aurantiacum  (LIGHT.)  NAEG.  and  HEPP.,  G. 

Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP.,  G. 

Lecanora  subfusca  (L.)  Acn.,  G. 

Pertusaria  pustulata  (Acn.)  NYL. 

Pertusaria  leioplaca  (Acn.)  SCHAKR. 

Pertusaria  velata  (TURN.)  NYL. 

Biatora  rubella  (EHRH.)  RABENH. 

Biatora  fuscorubella  (HOFFM.)  TUCK.,  G. 

Biatora  subfusca  FR.,  G. 

Lecidea  enteroleuca  FR.,  G. 

Buellia  alboatra  (HOFFM.)  TH.  FR.,  G. 

Buellia  parasema  (Acn.)  TH.  FR. 

Opegrapha  varia  (PERS.)  FR.,  G. 

Graphis  scripta  (L.)  ACH.,  G. 

Graphis  scripta  (L.)  ACH.  var.  limitata  ACH.,  G. 

Arthonia  lecideella  NYL. 

Arthonia  radiata  (PERS.)  TH.  FR.,  G. 

Coniocybe  pallida  (PERS.)  FR. 

Pyrenula  gemmata  (Acn.)  NAEG.,  G. 

Pyrenula  hyalospora  NYL.,  G. 

Pyrenula  nitida  ACH. 

Pyrenula  quinqueseptata  (NYL.)  TUCK. 

Pyrenula  leucoplaca  (WALLR.)  KBR.,  G. 

Pyrenula  megalospora  sp.  nov.,  G. 

In  order  to  avoid  reproducing  a  large  portion  of  the  above 
long  list  of  names,  I  have  for  the  similar  formation  at  Granite 
Falls  marked  those  of  the  list  found  there  (G)  and  add  below  the 


304  MINNESOTA    BOTANICAL    STUDIES. 

only  one  found  in  the  Granite  Falls  formation  and  not  at 
Mankato,  viz. ,  Bialora  naegelii  Tuck.  Thus  the  mark  (G)  will 
indicate  also  those  common  to  both  formations  and  as  a  whole 
most  characteristic  of  such  lichen  formations  for  the  Minnesota 
valley.  The  Mankato  area  with  its  abundance  of  trees  would, 
of  course,  be  expected  to  possess  richer  tree  lichen  formations 
than  Granite  Falls,  and  with  the  exception  of  a  single  species, 
the  rough  bark  formation  of  the  latter  area  is  but  a  partial  repe- 
tition of  that  of  the  former. 

The  formation  on  trees  with  smooth  bark  at  Mankato  con- 
tains all  but  two  of  the  species  of  the  similar  formation  at 
Granite  Falls,  and  the  treatment  may  be  abbreviated  as  the  last 
two  above.  The  genus  Pyrenula  predominates  in  the  forma- 
tion, and  some  of  the  species  are  among  the  lichens  most  char- 
acteristic of  smooth  bark.  Therefore,  the  formations  may  re- 
ceive the  name  which  follows  : 

Pyrenula  lichen  formation  of  trees  with  smooth  bark 
(Mankato). 

Theloschistes  polycarpus  (EHRH.)  TUCK. 

Theloschistes  concolor  (DICKS.)  TUCK.,  G. 

Parmelia  olivacea  (L.)  ACH.,  G. 

Physcia  adglutinata  (FLOERK.)  NYL.,  G. 

Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP.,  G. 

Lecanora subf usca  (L.)  ACH.,  G. 

Rinodina  sophodes  (AcH.)  NYL.,  G. 

Biatora  fuscorubella  (HOFFM.)  TUCK.,  G. 

Lecidea  enteroleuca  FR.,  G. 

Graphis  scripta  (L.)  ACH.,  G. 

Arthonia  lecideella  NYL. 

Arthonia  dispersa  NYL.,  G. 

Pyrenula  punctiformis  (Acn.)  NAEG.,  F. 

Pyrenula  punctiformis  (Acn.)  NAEG.  var.  fallax  NYL.,  F. 

Pyrenula  nitida  ACH.,  F. 

Pyrenula  thelena  ACH.,  F. 

Pyrenula  cinerella  (FLOT.)  TUCK.,  F. 

Pyrenula  cinerella  (FLOT.)  TUCK.  var.  quadriloculata  var.  nov. 

Pyrenula  leucoplaca  (WALLR.)  KBR.,  GF. 


Fink :   LICHENS  OF  THE  MINNESOTA  VALLEY.  305 

The  two  formed  on  smooth  bark  at  Granite  Falls  and  not  at 
Mankato  are  Lecidea  cntcroleuca  Fr.  var.  achrista  Schaer.  and 
Arthonia  punctiforniis  Ach.  As  in  the  rough  bark  formations, 
the  one  at  Mankato  is  richer  for  the  same  reason  and,  strangely 
enough,  my  study  of  the  Granite  Falls  area  only  discovered  a 
single  Pyrcnula  on  smooth  bark.  My  name  is  scarcely  appro- 
priate for  this  formation,  though  it  is  for  the  one  at  Mankato  as 
it  would  be  for  others  from  other  localities  in  Minnesota  and 
Iowa  which  might  be  added.  Without  adding  another  list  or 
another  complete  formation,  I  have  indicated  by  (F)  in  the 
above  list  the  Pyrenulas  of  that  list  which  occur  on  smooth 
bark  at  Fayette,  Iowa. 

Persons  acquainted  with  lichen  species  will  readily  observe 
in  the  lists  for  rough  bark  and  smooth  bark  lichen  formations, 
that  the  formation  on  rough  bark  is  composed  principally  of  the 
more  foliaceous  and  fruticulose  lichens  while  those  of  the  smooth 
bark  formations  are  in  the  main  crustaceous  lichens.  This  is 
possibly  due  in  part  to  the  fact  that  these  foliaceous  and  fruti- 
culose lichens  more  easily  gain  a  foothold  on  the  rough  bark 
which  breaks  up  the  thallus  of  the  lichens  adapted  to  smooth 
bark,  thus  tending  to  kill  them.  However  it  is  probable  that 
light,  shade  and  moisture  are  also  factors,  the  large  trees  fur- 
nishing more  shade  than  the  smaller  ones. 

Next  in  order  naturally  enough  we  may  consider  the  lichen 
formations  of  old  boards  and  old  wood,  and  the  formations  are 
so  nearly  alike  for  Mankato  and  Granite  Falls  that  they  may  be 
treated  in  a  single  list  by  giving  the  Mankato  list  and  marking 
(G)  those  common  to  the  Granite  Falls  formation  also.  Our 
Calicei  are  lichens  seldom  seen  in  any  other  formations,  hence 
the  following  name  may  be  applied. 

Calicei  lichen  formation  of  old  boards  and   wood 
(Mankato). 

'heloschistes   chrysopthalmus  (L.)  NORM.,  G. 

lacodium  cerinum  (HEDW).  NAEG.  and  HEPP.  var.  pyrocea 
NYL.,  G. 

Lecanora  hageni  ACH.,  G. 

Lecanora  varia  (EHRH.)  NYL.,  G. 

Rinodina  sophodes  (AcH.)  NYL.,  G. 

Rinodina  sophodes  (Acn.)  NYL.,  var.  exigua  FR.,  G. 


306  MINNESOTA    BOTANICAL    STUDIES. 

Buellia  parasema  (Acn.)  TH.  FR.,  G. 

Buellia  turgescens  (NYL.)  TUCK. 

Calicium  parietinum  ACH. 

Thelocarpon  prasinellum  NYL. 

The  additions  for  Granite  Falls  are  Cctraria  ciliaris  (Ach.) 
Tuck.,  Lecidea  entcroleuca  Fr.  and  Calicium  quercinum  Ach. 
As  in  other  instances  the  common  forms  are  those  most  charac- 
teristic of  such  formations.  I  have  not  detected  the  Calicium 
for  which  I  have  named  the  Mankato  formations  at  Granite 
Falls,  where  it  is  replaced  by  another  species,  and  I  shall  add 
the  species,  Acolium  tigillare  (Ach.)  Dn.,  which  is  one  of  the 
Calicei  common  in  the  similar  formation  at  Fayette,  Iowa,  and 
the  only  one  found  in  the  like  formation  at  Minneapolis.  It 
must  be  admitted  that  the  name  used  for  these  formations,  while 
it  may  be  applied,  is  not  so  appropriate  for  the  related  forma- 
tions in  the  lake  Superior  region  where  some  of  the  Calicei 
grow  on  living  bark  and  yet  others  on  rotting  wood. 

But  one  formation  remains  to  be  considered,  viz.,  that  of  rot- 
ting stumps  and  prostrate  logs.  In  these  formations  the  most 
common  plants  are  those  of  the  genus  Cladonia  and  the  forma- 
tions may  accordingly  receive  the  following  name  : 

Cladonia  formation  of  rotten  2uood  (Mankato). 
Peltigera  canina  (L.)  HOFFM.,  G. 
Peltigera  canina  (L.)  HOFFM.,  var.  sorediata  SCHAER. 
Cladonia  fimbriata  (L.)  FR.,  G. 
Cladonia  fimbriata  (L.)  FR.  var.  tubaBformis  FR.,  G. 
Cladonia  gracilis  (L.)  NYL.,  G. 
Cladonia  gracilis  (L.)  NYL.,  var.  verticiilata  FR.,  G. 
Cladonia  symphycarpia  TUCK. 
Cladonia  macilenta  (EHRH.)  HOFFM. 
Cladonia  cristatella  TUCK. 

The  only  species  found  at  Granite  Falls  in  the  similar  forma- 
tion and  not  at  Mankato  is  Biatora  flexuosa  Fr.  and  the  forma- 
tion may,  with  this  addition,  be  indicated  by  marking  (G)  those 
plants  of  the  Mankato  formation  common  to  both.  Comparison 
with  formations  from  other  localities  would  show  some  varia- 
tion, but  the  Cladonias  would  predominate  and  give  character 
to  the  formations.  Wood  commonly  rots  in  moist  shady  places. 


Fink  :     LICHENS    OF    THE    MINNESOTA    VALLEY. 

furnishing  an  abundance  of  moisture,  and  we  find  accordingly 
that  the  formations  on  rotten  wood  are  made  up  in  large  part  of 
fruticulose  lichens.  The  Calicei  formations  of  old  wood  are 
exposed  to  drier  conditions  and  are  composed  almost  entirely  of 
lichens  having  poorly  developed  thalli. 

I  must  emphasize  here  that  lichens  of  nearly  all  the  forma- 
tions enumerated  above  enjoy  moist  places,  and  that  lack  of 
moisture  produces  a  decrease  in  richness  both  in  size  and  num- 
ber of  individuals  and  in  numbers  of  species  in  the  formations. 
I  repeat  this,  which  I  have  established  for  some  parts  of  Minne- 
sota previously,  because  some  persons  may  suppose  that  lichens, 
because  of  their  xerophytic  adaptations,  thrive  as  well  in  the 
driest  spots  as  in  those  affording  more  moisture.  The  excep- 
tions to  this  general  statement  will  appear  from  a  careful  study 
of  the  analyses  made  of  the  various  formations. 

The  gelatinous  lichen  formation  of  shaded  limestone  (Minne- 
apolis) has  been  called  a  scattered  one,  and  I  have  explained 
what  is  meant  by  the  expression.  Others  of  the  same  kind  are 
the  Cladonia-Peltigera  lichen  formations  of  shaded  earth,  the 
angiocarpous  lichen  formations  of  exposed  limestone  bluffs,  the 
Calicei  lichen  formations  of  old  wood  and  in  some  instances  the 
Cladonia  lichen  formations  of  rotten  wood,  though  in  other  in- 
stances half  or  more  of  the  species  of  Cladonia  of  the  forma- 
tion may  be  found  on  a  single  log.  Thus  formations  of  the 
kind  last  named  and  like  the  one  first  named  in  this  paragraph 
differs  from  the  other  three  named  in  the  paragraph  in  that  they 
may  or  may  not  be  scattered  while  the  three  always  are,  so  far 
as  I  know,  except  the  Calicei  formation  which  may  be  found 
nearly  complete  on  a  few  rods  of  old  fence  in  some  favorable 
instances.  The  two  formations  of  trees  are  widely  extended  ; 
but  they  are  not  scattered  as  I  have  used  the  term  since  one 
commonly  finds  a  good  proportion  of  the  species  of  either  for- 
mation in  passing  a  short  distance,  in  the  woods. 

Also  in  my  classification  we  have  the  peculiar  condition  of 
two  lichen  formations  occupying  the  same  area.  This  is  illus- 
trated by  the  Biatora  decipiens  lichen  formation  of  exposed 
calcareous  earth  and  the  Lecanora  calcarea  contorta  lichen 
formation  of  exposed  limestone  pebbles,  or  by  the  Lecanora 
lichen  formations  of  exposed  granite  or  quartzite  and  the  Endo- 
carpon  hepaticum  lichen  formation  of  exposed  earth.  Yet  it  is 
apparent  that  the  formations  are  distinct  in  both  instances,  the 


308  MINNESOTA    BOTANICAL    STUDIES. 

division  being  based  on  substratum  as  well  as  amount  of  light 
and  moisture. 

As  a  whole,  the  formations  may  be  said  to  be  azonal  and 
without  definite  form  or  extent,  both  depending  upon  location 
of  proper  substrata,  protection  from  or  exposure  to  light,  etc. 

In  my  paper  I  have  used  the  expression  "  lichen  formation" 
to  include  lichens  only.  Of  course,  these  plants  are  in  some 
instances  found  growing  upon  the  same  substrata  and  in  the 
same  general  set  of  conditions  as  plants  of  other  groups,  and 
which  might  have  been  listed  in  the  formations.  However,  I 
may  be  excused,  in  a  paper  on  lichen  distribution,  for  omitting 
other  plants  than  lichens,  especially  since  I  could  not  possibly 
have  treated  the  other  plants  with  the  same  detail  that  I  have 
accorded  the  lichens. 

I  know  of  no  other  paper  which  has  dealt  exclusively  with 
lichen  distribution  as  I  have  done  herein,  and  surely  this  anal- 
ysis must  be  helpful  in  the  study  of  the  lichen  flora  of  other 
regions.  The  multiplicity  of  observations  necessary  for  such  a 
detailed  study  are  not  easy  to  make,  and  I  am  sure  that  much 
of  interest  has  escaped  me.  However,  I  hope  that  this  paper 
may  stimulate  others  to  study  the  lichens  from  an  ecologic  point 
of  view. 

LIST    OF    SPECIES    AND    VARIETIES. 

1.  Ramalina  calicaris  (L.)  FR.  var.  fraxinea  FR. 

On  trees  and  old  wood,  infrequent  or  rare.  Mankato,  June 
23,  1899,  no.  55,  June  26,  1899,  no.  102,  and  June  28,  1899, 
no.  164.  Granite  Falls,  July  n,  1899,  no.  385  and  July  13, 
1899,  nos.  510  and  533. 

2.  Ramalina  calicaris  (L.)  FR.  var.  fastigiata  FR. 

On  trees  and  rocks,  rare.  Mankato,  June  23,  1899,  no.  54. 
New  Ulm,  July  5,  1899,  no.  275.  Granite  Falls,  July  14,  1899, 
no.  518,  and  July  17,  1899,  no.  588. 

3.  Ramalina  calicaris  (L.)  FR.  var.  farinacea  SCHAER. 

On  sandstone  and  granite.  Mankato  ( Minneopa  Falls) ,  June 
27,  1899,  no.  154.  Redwood  Falls,  July  6,  1899,  no.  305,  and 
July  8,  1899,  no.  349.  Granite  Falls,  July  12,  1899,  no.  460. 

4.  Ramalina  polymorpha  (Acn.)  TUCK.  ? 

On  shaded  granitic  rocks  in  first  locality  and  on  a  large 
bowlder  in  the  second,  rare.  Granite  Falls,  July  12,  1899,  no. 


Fink :   LICHENS  OF  THE  MINNESOTA  VALLEY.  309 

456,  and  July  13,  1899,  no.  492.  Pipestone,  July  19,  1899,  no. 
641. 

The  plants  are  placed  here  provisionally. 

They  resemble  in  part  Ramalina  calicaris  (L.)  Fr.  \-&x.  fari- 
nacea  Schaer; 

Not  previously  reported  from  Minnesota  and  new  to  the  Mis- 
sissippi valley. 

5.  Cetraria  ciliaris  (Acn.)  TUCK. 

A  single  sterile  specimen  collected  on  an  old  cedar  stump. 
Granite  Falls,  July  17,  1899,  no.  570. 

6.  Usnea  barbata  (L.)  FR.  var.  florida  FR. 

On  an  old  stump,  only  seen  once  and  then  sterile.  Granite 
Falls,  July  17,  1899,  no-  5^5 • 

7.  Usnea  barbata  (L.)  FR.  var.  hirta  FR. 

On  sandstone,  rare.  Mankato  (Minneopa),  June  27,  1899, 
no.  151. 

8.  Usnea  barbata  (L.)  FR.  var.  rubiginea  MICHX. 

On  sandstone  and  granite  rocks,  rare.  Mankato  (Minneopa 
Falls),  June  27,  1899,  no.  152.  Redwood  Falls,  July  8,  1899, 
no.  350. 

9.  Theloschistes  chrysopthalmus  (L.)  NORM. 

On  trees  and  old  boards,  rare  or  infrequent  except  at  Granite 
Falls,  where  the  plant  is  frequent.  Mankato,  June  22,  1899, 
no.  9.  Mankato  (Rapidan),  June  28,  1899,  no.  163.  New 
Ulm,  July  4,  1899,  nos.  226  and  227.  Redwood  Falls,  July  6, 
1899,  no.  302,  and  July  8,  1899,  no.  340.  Granite  Falls,  July 
u,  1899,  no.  387,  and  July  15,  1899,  no-  549- 

10.  Theloschistes  polycarpus  (EHRH.)  TUCK. 

On  trees  and  rocks,  rare.  Mankato,  June  26,  1899,  no.  107. 
Granite  Falls,  July  12,  1899,  no.  447,  and  July  15,  1899,  no.  531. 

u.  Theloschistes  lychneus  (NYL.)  TUCK. 

On  trees  and  rocks,  frequent.  Mankato,  June  22,  1899,  no. 
5.  New  Ulm,  July  5,  1899,  no.  263.  Redwood  Falls,  July  8, 
1899,  no.  330.  Granite  Falls,  July  11,  1899,  no.  438,  and  July 
17,  1899,  no.  579. 

12.  Theloschistes  concolor  (DICKS.)  TUCK. 

On  trees  and  old  wood,  common  at  the  first  locality  and  rare 
at  the  second.  Mankato,  June  22,  1899,  no.  7.  Granite  Falls, 
July  n,  1899,  no.  377. 


310  MINNESOTA    BOTANICAL    STUDIES. 

13.  Theloschistes  concolor  (DICKS.)  TUCK.  var.  effusa  TUCK. 
On  trees  rare.     Mankato,  July  i,  1899,  no.  2i6a. 

14.  Parmelia  perforata  (JACQ^)  ACH. 

On  trees,  rare.      Mankato,  June  26,  1899,  no.   134. 

15.  Parmelia  cetrata  ACH. 

On  trees  and  rocks,  rare  except  at  the  last  locality  where  the 
plant  is  frequent.  New  Ulm,  July  4,  1899,  no.  228.  Redwood 
Falls,  July  8,  1899,  nos.  329  and  360.  Granite  Falls,  July  12, 
1899,  no.  455,  and  July  17,  1899,  nos.  550,  573  and  574. 

1 6.  Parmelia  crinita  ACH. 

On  trees  and  granitic  rocks,  rare.  Mankato,  June  23,  1899, 
no.  47.  Mankato  (Minneopa  Falls),  June  27,  1899,  no.  133. 
New  Ulm,  July  5,  1899,  no.  258.  Granite  Falls,  July  n, 
1899,  nos.  400  and  439. 

17.  Parmelia  borreri  TURN. 

On  trees  and  granitic  rocks,  common.  Mankato,  June  22, 
1899,  no.  ii.  New  Ulm,  July  5,  1899,  no-  2^8.  Redwood 
Falls,  July  8,  1899,  no.  335.  Granite  Falls,  July  n,  1899,  nos. 
383  and  389. 

18.  Parmelia  borreri  TURN.  var.  rudecta  TUCK. 

On  trees  and  old  wood,  infrequent.  Mankato,  June  23,  1899, 
no.  12,  and  July  23,  1899,  no.  58. 

19.  Parmelia  borreri  TURN.  var.  hypomela  TUCK. 

On  shaded  granite  rocks,  rare  and  sterile.  New  Ulm,  July 
5,  1899,  no.  287. 

20.  Parmelia  tiliacea  (HOFFM.)  FLOERK. 

On  trees,  infrequent.     Mankato,  June  23,  1899. 

21.  Parmelia  saxatilis  (L.)  FR. 

On  trees  and  rocks,  rare.  Mankato,  June  26,  1899,  no-  IO^- 
New  Ulm,  July  5,  1899,  no-  2^4-  Redwood  Falls,  July  8, 
1899,  no-  341-  Granite  Falls,  July  n,  1899,  no.  414. 

22.  Parmelia  saxatilis  (L.)  FR.  var.  sulcata  NYL. 

On  old  wood  and  shaded  rocks,  rare.  Mankato  (Rapidan), 
June  28,  1899,  no.  165.  Granite  Falls,  July  17,  1899,  no.  586. 

23.  Parmelia    saxatilis     (L.)    FR.    var.    panniformis    (Acn. 

(SCHAER. ? 

On   shaded   rocks,  rare.     New   Ulm,  July  5,  1899,  no.  268. 
Granite  Falls,  July  17,  1899,  no  561. 
The  plant  is  placed  here  provisionally. 


Fink  :     LICHENS    OF    THE    MINNESOTA    VALLEY.  311 

Not  previously  reported  from  Minnesota  and  new  to  the 
Mississippi  valley. 

24.  Parmelia  olivacea  (L.)  Ach. 

On  trees  and  old  wood,  rare.  Mankato,  June  23,  1899,  no. 
61.  Granite  Falls,  July  15,  1899,  no-  537- 

25.  Parmelia  olivacea  (L.)  ACH.  var.  prolixa  ACH. 

On  granitic  rocks,  quartzite,  pipestone  and  once  collected  on 
earth,  frequent  except  at  the  first  locality,  where  it  is  rare. 
Morton,  July  7,  1899,  no.  315.  Granite  Falls,  July  n,  1899, 
no.  405.  Pipestone,  July  18,  1899,  nos.  594,  609  and  621,  and 
July  19,  1899,  no.  643. 

26.  Parmelia  caperata  (L.)  ACH. 

On  trees  and  granitic  rocks,  frequent.  Mankato,  June  23, 
1899,  no.  53.  Mankato  (Minneopa)  June  27,  1899,  no.  153. 
New  Ulm,  July  5,  1899,  no.  285.  Granite  Falls,  July  u, 
1899,  no-  441- 

27.  Parmelia  conspersa  (£HRH.)  ACH. 

On  granitic  rocks,  quartzite  and  pipestone,  common  or  abun- 
dant. New  Ulm,  July  5,  1899,  no.  269.  Granite  Falls,  July 
12,  1899,  no.  449.  Pipestone,  July  18,  1899,  no-  5^9»  an^ 
July  19,  1899,  no.  637. 

28.  Physcia  speciosa  (WULF.,  ACH.)  NYL. 

On  rocks  and  mossy  bases  of  trees,  infrequent.  Mankato, 
June  22,  1899,  no.  13.  New  Ulm.,  July  5,  1899,  no.  281. 
Granite  Falls,  July  u,  1899,  no-  374- 

29.  Physcia  granulifera  (Acn.)  TUCK. 

On  trees,  rare.  Mankato,  June  24,  1899,  no.  84.  Granite 
Falls,  July  n,  1899,  no.  372. 

30.  Physcia  pulverulenta  (SCHREB.)  NYL. 

On  rocks  and  trees,  frequent.  Mankato,  June  23,  1899,  no. 
52.  New  Ulm,  July  5,  1899,  no.  286.  Granite  Falls,  July  u, 
1899.  no.  383. 

31.  Physcia  stellaris  (L.)  TUCK. 

On  trees  and  rocks,  common  or  abundant.  Mankato,  June 
23,  1899,  no-  I-  New  Ulm,  July  5,  1899,  no  297.  Granite 
Falls,  July  u,  1899  no.  431,  and  July  13,  1899,  no.  481. 

32.  Physcia  stellaris  (L.)  TUCK.  var.  apiola  NYL. 

On  granitic  rocks,  infrequent.  Mankato,  June  23,  1899,  no. 
44.  Granite  Falls,  July  12,  1899,  no.  463. 


312  MINNESOTA    BOTANICAL    STUDIES. 

33.  Physcia  tribacia  (Acn.)  TUCK. 

On  wood,  granite  and  quartzite,  rare.  Mankato,  June  23 ,. 
1899,  no.  77,  and  July  i,  1899,  no.  216.  Granite  Falls,  July 
14,  1899,  no.  517.  Pipestone,  July  18,  1899,  no.  601  and  July 
19,  1899,  no.  634. 

34.  Physcia  caesia  (HOFFM.)  NYL. 

On  bowlders  and  all  kinds  of  rocks  in  the  region,  except  lime- 
stone, frequent.  Mankato,  June  23,  1899,  no.  76.  New  Ulm, 
July  5,  1899,  nos.  290  and  296.  Granite  Falls,  July  12,  1899, 
no.  443.  Pipestone,  July  18,  1899,  nos.  618  and  624. 

35.  Physcia  obscura  (EHRH.)  NYL. 

On  trees  and  rocks,  common.  Mankato,  June  1899,  no-  7^a- 
New  Ulm,  July  5,  1899,  no.  289.  Granite  Falls,  July  n,  1899, 
no.  378,  and  July  17,  1899,  no.  583. 

36.  Physcia  adglutinata  (FLOERK.)  NYL. 

On  trees,  frequent.  Mankato,  June  23,  1899,  no.  45.  Gran- 
ite Falls,  July  n,  1899,  no.  382  and  July  13,  1899,  no.  482. 

37.  Pyxine  sorediata  FR. 

On  granitic  rocks,  rare.  Granite  Falls,  July  17,  1899,  no. 
578. 

38.  Peltigera  rufescens  (NECK.)  HOFFM. 

On  earth  and  mossy  rocks,  frequent.  Mankato,  June  25,. 
1899,  no.  48.  Mankato  (Minneopa  Falls),  June  27,  1899,  no. 
150.  New  Ulm,  July  5,  1899,  no.  251.  Granite  Falls,  July 
n,  1899,  no.  395  and  July  17,  1899,  no.  559. 

39.  Peltigera  canina  (L.)  HOFFM. 

On  earth  and  rocks,  common.  Mankato  (Minneopa  Falls),. 
June  27,  1899,  no.  149.  New  Ulm,  July  5,  1899,  no.  262. 
Granite  Falls,  July  n,  1899,  no.  390,  and  July  17,  1899,  no- 
580. 

40.  Peltigera  canina  (L.)  HOFFM.  var.  spongiosa  TUCK. 
On  earth,  rare.      Redwood  Falls,  July  8,  1899,  no.  357. 
Not  previously  reported  from  Minnesota. 

41.  Peltigera  canina  (L.)  HOFFM.  var.  spuria  ACH. 

On  earth,  rare.     Mankato  (Rapidan),  June  28,  1899,  no.  161. 

42.  Peltigera  canina  (L.)  HOFFM.  var.  sorediata  SCHAER. 

On  earth  and  old  wood,  rare,  Mankato,  June  26,  1899,  no. 
121.  New  Ulm,  July  5,  1899,  no.  260.  Granite  Falls,  July 
13,  1899,  no.  512. 


Fink  :    LICHENS  OF  THE  MINNESOTA  VALLEY.  313 

43.  Peltigera  horizontalis  (L.)  HOFFM. 

On  shaded  earth,  frequent  locally.  Redwood  Falls,  July  6, 
1899,  no.  301. 

44.  Heppia  despreauxii  (MONT.)  TUCK. 

On  earth,  rare  at  first  locality  and  frequent  at  second.  Man- 
kato  (Rapidan)  June  28,  1899.  no.  177.  Granite  Falls,  July  n, 
1899,  no.  394,  July  13,  1899,  no.  507  and  July  14,  1899,  no. 
522. 

Not  previously  reported  from  Minnesota. 

45.  Heppia  polyspora  TUCK.  ? 

On    earth,    rare.      Granite    Falls,  July    13,    1899,   no.    498. 

<— 8 
Spores  spherical   or  subspherical,       d     mic.      This      exceeds 

5     . 

Tuckerman's  measurements.  Apothecia  occasionally  surpass- 
ing one  mm.  in  diameter.  I  may  later  find  it  necessary  to 
separate  this  as  new  species. 

Not  previously  reported  from  Minnesota. 

46.  Pannaria  languinosa  (Acn.)  KOERB. 

On  various  rocks,  earth  and  trees  in  shaded  places,  common 
or  abundant.  Mankato,  June  23,  1899,  no.  50.  New  Ulm, 
July  5,  1899,  no.  280.  Granite  Falls,  July  n,  1899,  no.  396, 
and  July  17,  1899,  no.  569. 

47.  Pannaria  microphylla  (Sw.)  DELIS. 

On  shaded  granite,  infrequent.  Redwood  Falls,  July  8, 
1899,  no.  345.  Granite  Falls,  July  u,  1899,  no.  384. 

48.  Pannaria  nigra  (Huos.)  NYL. 

On  limestone,  common  locally.  Mankato,  June  24,  1899, 
no.  95. 

49.  Ephebe  pubescens  FR. 

On  quartzite,  rare.     New  Ulm,  July  5,  1899,  no.  257. 

50.  Pyrenopsis  phaeococca  TUCK. 

On  bowlders,  rare.     Mankato,  June  23,  1899,  no.  74. 
Not    previously   reported    from    Minnesota    and   new   to    the 
Mississippi  valley. 

51.  Pyrenopsis  melambola  TUCK.? 

On  bowlders,  frequent.  Mankato,  June  29,  1899,  no.  189. 
Spores  somewhat  small  (7-10x4-5*^  mic.). 

Not  previously  reported  from  Minnesota  and  new  to  the 
Mississippi  valley. 


314  MINNESOTA    BOTANICAL    STUDIES. 

52.  Omphalaria  kansana  TUCK. 

On  limestone,  locally  frequent.  Mankato,  June  23,  1899, 
no.  27. 

Not  previously  reported  from  Minnesota. 

53.  Omphalaria  pulvinata  NYL. 

On  limestone,  rare.     Mankato,  June  27,  1899,  no.  148. 
Not  previously  reported  from  Minnesota. 

54.  Omphalaria  phyllisca  (WAHL.)  TUCK. 

On  granitic  rocks,  rare.  Redwood  Falls,  July  6,  1899,  no- 
360,  and  July  8,  1899,  nos.  338  and  343.  Granite  Falls,  July 
17,  1899,  nos.  572  and  584. 

Not  previously  reported  from  Minnesota  and  new  to  the 
Mississippi  valley. 

55.  Collema  pycnocarpum  NYL. 

On  trees  and  once  on  rocks,  generally  distributed  in  the 
Minnesota  valley,  but  rare. 

Mankato,  June  23,  1899,  no.  60,  and  June  24,  1899,  nos.  89 
and  97.  Mankato  (Minneopa  Falls),  June  27,  1899,  no.  136. 
Redwood  Falls,  July  6,  1899,  no.  309,  and  July  8,  1899,  no. 
355.  Granite  Falls,  July  u,  1899.  no.  380. 

56.  Collema  flaccidum  ACH. 

On  trees  and  rocks,  rare.  Mankato,  June  23,  1899,  no.  81. 
Mankato  (Minneopa),  June  27,  1899,  no.  137.  New  Ulm,  July 
5,  1899,  no.  278. 

57.  Collema  pulposum  (.BERNH.)  NYL. 

On  earth  and  rocks,  common  in  first  locality.  Mankato, 
June  22,  1897,  no.  3,  and  June  23,  1899,  no.  79.  Granite 
Falls,  July  u,  1899,  no.  417. 

58.  Collema  tenax  (Sw.)  ACH. 

On  earth,  rare.     Mankato  (Rapidan),  June  28,  1899,  no.  169. 
Not  previously  reported  from  Minnesota. 

59.  Collema  plicatile  SCHAER. 

On  calcareous  rocks,  locally  frequent.  Mankato,  June  23, 
1899,  no.  28. 

Not  previously  reported   from   Minnesota. 

60.  Collema  pustulatum  ACH. 

On  calcareous  rocks,  rare.  Mankato,  June  26,  1899,  no' 
in. 

Not  previously  reported  from  Minnesota. 


li"ink  :    LiriiKxs  OF  THE  MINNESOTA  VALLEY. 

61.  Collema  furvum  (Acn.)  NYL. 

On  shaded  rocks,  infrequent.  New  Ulm,  July  5,  1899,  no- 
283.  Redwood  Falls,  July  6,  1899,  no.  307.  Granite  Falls, 
July  u,  1899,  no-  391- 

Not  previously  reported  from  Minnesota. 

62.  Leptogium  lacerum  (Sw.)  FR. 

On  various  rocks,  usually  shaded,  frequent.  Mankato,  June 
23,  1899,  no.  49.  Redwood  Falls,  July  8,  1899,  no.  328. 
Granite  Falls,  July  13,  1899,  nos.  480  and  497  and  July  17, 
1899,  no.  568. 

63.  Leptogium  pulchellum  (Acn.)  NYL. 

Collected  once  on  a  large  bowlder  in  a  shaded  ravine,  rare. 
Mankato,  July  i,  1899,  no.  212. 

Not  previously  reported  from  Minnesota. 

64.  Leptogium  chloromelum  (Sw.)  NYL. 

On  mossy,  shaded  sandstone,  rare.  Mankato  (Minneopa 
Falls),  June  27,  1899,  no.  144. 

65.  Leptogium  myochroum  (EHRH.,  SCHAER.)  TUCK. 

On  trees  and  shaded  granitic  rocks,  rare.  Mankato,  June 
26,  1899,  no.  126.  Mankato  (Rapidan),  June  28,  1899,  no. 
166.  Granite  Falls,  July  n,  1899,  no.  392. 

66.  Placodium  elegans  (LINK.)  DC. 

On  various  rocks  ;  common  at  Granite  Falls,  infrequent  else- 
where. Mankato,  June  24,  1899,  no.  83.  Redwood  Falls, 
July  8,  1899,  no.  353.  Granite  Falls,  July  11,  1899,  no.  440. 
Pipestone,  July  18,  1899,  nos-  592  and  603. 

67.  Placodium  murorum  (HOFFM.)  DC. 

On  granitic  rocks,  rare.  Granite  Falls,  July  12,  1899,  no. 
452. 

68.  Placodium  cinnabarrinum  (Acn.)  Auz. 

On  various  rocks,  frequent  or  common.  Mankato,  June  23, 
1899,  no-  73»  and  June  30,  1899,  no.  200.  Morton,  July  7, 
1899,  no.  320.  North  Redwood,  July  10,  1899,  no.  369.  Gran- 
ite Falls  July  n,  1899,  no>  411-  Pipestone,  July  18,  1899, 
no.  607. 

69.  Placodium  aurantiacum  (LIGHTF.)  NAEG.  and  HEPP. 

On  trees  and  rocks,  common  at  first  locality.  Mankato, 
June  22,  1899,  no.  19,  and  June  23,  1899.  no.  38.  Granite 
Falls,  July  n,  1899,  no.  375. 


316  MINNESOTA    BOTANICAL    STUDIES. 

70.  Placodium  citrinum  (HOFFM.)  LEIGHT. 

On  limestone,  rare.      Mankato,  June  26,  1899,  no-  II7- 

71.  Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP. 

On  trees  and  old  wood,  common.  Mankato,  June  22,  1899,. 
no.  18,  and  June  23,  1899,  no.  75.  Mankato  (Rapidan),  June 
28,  1899,  nos.  178  and  179.  Granite  Falls,  July  n,  1899,  nos. 
432  and  435.  Granite  Falls,  July  17,  1899,  no.  553. 

72.  Placodium  cerinum  (HEDW.)  NAEG.  and   HEPP.  var.  sider- 
itis  TUCK. 

On  granitic  rocks  and  catlinite,  common.  Mankato,  June 
23,  1899,  no.  41,  June  26,  1899,  no.  114,  and  June  30,  1899, 
no.  198.  New  Ulm,  July  5,  1899,  no.  247.  Granite  Falls, 
July  12,  1899,  no.  444.  Pipestone,  July  18,  1899,  no.  613. 

73.  Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP.  var.  pyracea 
NYL. 

On  old  boards,  infrequent.  Mankato,  June  22,  1899,  no.  2. 
Granite  Falls,  July  n,  1899,  no.  430. 

74.  Placodium  ferrugineum  (Huos.)  HEPP. 

On  old  wood,  rare.     Mankato,  June  22,  1899,  no.  17. 
Not  previously  reported  from  Minnesota. 

75.  Placodium  ferrugineum  (Huos.)  HEPP.  var.  pollinii  TUCK. 
On  dead  cedars,  rare.     New  Ulm,  July  4,  1899,  no.  230. 
Not  previously  reported  from  Minnesota. 

76.  Placodium  vitellinum  (EHRH.)  NAEG.  and  HEPP. 

On  granite  and  quartzite,  common.  Mankato,  June  30, 
1899,  no.  203.  New  Ulm,  July  5,  1899,  no.  292.  Granite 
Falls,  July  12,  1899,  no-  4^2>  Pipestone,  July  18,  1899,  no- 
629. 

77.  Placodium  vitellinum  (EHRH.)  NAEG.  and  HEPP.  var.  aurel- 
lum  ACH. 

On  granite,  quartzite  and  sandstone,  frequent.  Mankato, 
June  22,  1899,  no.  10.  New  Ulm,  July  4,  1899,  nos.  233,  237 
and  239.  Granite  Falls,  July  14,  1899,  no.  523. 

78.  Lecanora  sp. 

On  granitic  rocks,  frequent  locally.      Granite  Falls,  July  n, 

1899,    no.    408.     Spores  -~ mic.      Seems    near   Lecanora 

x  "  3 

gelida  (L.)  Ach.,  but  the  thallus   is   probably  too   rough   and 
heavy. 


Fink:    LICHENS  OF  THE  MINNESOTA   VALLEY.  317 

Not  previously  reported  from  Minnesota  and  new  to  the  Miss- 
issippi Valley. 

79.  Lecanora  rubina  (Vn.L. )  Acn. 

On  granite,  quartzite  and  pipestone,  common.  Mankato 
(Rapidan),  June  28,  1899,  no.  160.  New  Ulm,  July  5,  1899, 
no.  248.  Granite  Falls,  July  n,  1899,  no.  401  and  July  12, 
1899,  no>  451-  Pipestone,  July  18,  1899,  no.  617,  and  July  19, 
1899,  no.  644. 

80.  Lecanora  rubina  (Vn,L.)  ACH.  var.  heteromorpha  ACH. 
With  the  last,  frequent.     New  Ulm,  July  5,  1899,    no.    266. 

North  Redwood,  July  10,  1899,  no.  361.  Granite  Falls,  July 
n,  1899,  no.  409.  Pipestone,  July  18,  1899,  and  July  19,  1899, 
no.  642. 

81.  Lecanora  muralis  (SCHREB.)  SCHAER. 

On  calcareous  rocks,  granite  and  quartzite,  common  at  Granite 
Falls,  rare  elsewhere.  Mankato,  June  24,  1899,  no.  85. 
Granite  Falls,  July  n,  1899,  no.  406.  Pipestone,  July  18, 
1899,  no.  630. 

82.  Lecanora  muralis  (SCHREB.)  SCHAER.  var.  versicolor  FR. 
On  calcareous  rocks,  rare.     Mankato,  June  30,  1899,  no.  196. 

83.  Lecanora  muralis  (SCHREB.)  SCHAER.  var.  saxicola  SCHAER. 
On   granitic   rocks   and   catlinite,  frequent.      Mankato,  June 

30,  1899,  no.  202.  North  Redwood,  July  10,  1899,  no.  370. 
Pipestone,  July  19,  1899,  no.  638. 

84.  Lecanora  frustulosa  (DICKS.)  MASS. 

On  rocks,  rare.  Redwood  Falls,  July  8,  1899,  no.  351. 
North  Redwood,  July  10,  1899,  no.  371.  Granite  Falls,  July 
n,  1899,  no.  410. 

85.  Lecanora  subfusca  (L.)  ACH. 

On  trees  and  rocks,  common  at  Mankato  only.  Mankato, 
June  23,  1899,  no.  43,  and  June  26,  1899,  no.  125.  New  Ulm, 
July  5,  1899,  no-  27Z-  Granite  Falls,  July  u,  1899,  no.  376. 

86.  Lecanora  subfusca  (L.)  ACH.  var.  allophana  ACH. 

On  granitic  rocks,  infrequent.  Granite  Falls,  July  n,  1899, 
nos.  403  and  407. 

Not  previously  reported  from  Minnesota  : 

87.  Lecanora  subfusca  (L.)  ACH.  var.  argentata  ACH. 
On  trees,  rare.     Mankato,  June  23,  1899,  no.  78- 


318  MINNESOTA    BOTANICAL.    STUDIES. 

88.  Lecanora  subfusca  (L.)  ACH.  var.  coilocarpa  ACH. 

On  trees,  granite  and  sandstone,  rare.  Mankato  (Rapidan), 
June  28,  1899,  no.  i6^a.  Mankato,  July  i,  1899,  no.  210. 
Granite  Falls,  July  12,  1899,  no.  446. 

89.  Lecanora  subfusca  (L.)  ACH.  var.  distans  ACH. 

On  sandstone,  rare.  Mankato  (Rapidan),  June  28,  1899,  no. 
182. 

90.  Lecanora  hageni  ACH. 

On  calcareous  and  granitic  rocks  and  on  old  boards,  common. 
Mankato,  June  21,  1899,  no  91,  and  July,  i,  1899,  nos.  215, 
217  and  218.  Granite  Falls,  July  u,  1899,  nos.  426  and  436, 
and  July  13,  1899,  no.  506. 

91.  Lecanora  varia  (EHRH.)  NYL. 

On  old  wood  and  trees,  infrequent.  Mankato,  June  24,  1899, 
no.  86.  Mankato  (Rapidan),  June  28,  1899,  no.  159.  New 
Ulm,  July  5,  1899,  no.  280.  Granite  l<alls,  July  n,  1899, 
no.  386. 

92.  Lecanora  erysibe  NYL. 

On  granitic  rocks,  rare.  Mankato,  June  23,  1899,  no.  63, 
and  June  26,  1899,  no.  131.  Granite  Falls,  July  13,  1899,  no. 

504- 

93.  Lecanora  cinerea  (L.)  SOMMERF. 

On  granite,  quartzite  and  catlinite,  common.  Mankato, 
June  6,  1899,  no.  62.  New  Ulm,  July  5,  1899,  no.  261.  Gran- 
ite Falls,  "July  13,  1899,  no.  493.  Pipestone,  July  18,  1899, 
nos.  625  and  633. 

94.  Lecanora  cinerea  (L.)  SOMMERF.  var.  laevata  FR. 
On  quartzite,  rare.     New  Ulm,  July  5,  1899,  no.  277. 

95.  Lecanora  cinerea  (L.)  SOMMERF.  var.  gibbosa  NYL. 
On  bowlders,  rare.     Mankato,  July  i,  1899,  no.  221. 

96.  Lecanora  calcarea  (L.)  SOMMERF. 

On  limestone,  rare.     Mankato,  June  29,  1899,  no.  188. 

97.  Lecanora  calcarea  (L.)  SOMMERF.  var.  contorta  FR. 

On  limestone,  drift  pebbles  and  granite,  infrequent.  Man- 
kato, June  30,  1899,  no.  199.  Redwood  Falls,  July  8,  1899, 
no.  333.  Granite  Falls,  July  n,  1899,  no.  402,  and  July  14, 
1899,  no.  524. 


Fink :   LICHENS  OF  THE  MINNESOTA  VALLEY.  319 

98.  Lecanora  xanthophana  NYL. 

On  granite,  quartzite  and  pipestone,  common.  Mankato, 
June  30,  1899,  no.  197.  New  Ulm,  July  5,  1899,  no.  246. 
Morton,  July  7,  1899,  no.  313.  Redwood  Falls,  July  8,  1899, 
no.  331.  North  Redwood,  July  10,  1899,  no.  368.  Granite 
Falls,  July  u,  1899,  no.  418.  Pipestone,  July  18,  1899,  nos- 
598  and  611. 

99.  Lecanora  cervina  (PERS.)  NYL. 

On  bowlders  and  sandstone,  infrequent.  Mankato  (Rapidan), 
June  28,  1899,  no.  179.  Mankato,  June  29,  1899,  no.  192. 

100.  Lecanora  cervina  (PERS.)  NYL.  var.  cinereoalba  var.  nov. 
On  granite,    frequent.     Mankato,   June  29,  1899,    no.    190. 

Granite    Falls,  July    u,  1899,  nos-    3^5  an^  4°3'  an^  Juty  I2» 
1899,  no.  404. 

Thallus  gray  or  grayish  white. 

101.  Lecanora  fuscata  (SCHRAD.)  TH.  FR. 

On  bowlders,  common  at  Mankato.  Mankato,  June  29,  1899, 
no.  191.  Granite  Falls,  July  12,  1899,  no.  450. 

102.  Lecanora  bookii  (FR.)  TH.  FR. 

On  limestone,  rare.     Mankato,  June  29,  1899,  no.  193. 
Not  previously  reported  from  Minnesota  and  new  to  the  Mis- 
sissippi valley. 

103.  Lecanora  privigna  (Acn.)  NYL. 

On  sandstone  and  calcareous  drift  pebbles,  rare.  Mankato 
(Rapidan),  June  28,  1899,  no.  171.  Granite  Falls,  July  13, 
1899,  no.  508. 

104.  Lecanora  privigna  (Acn.)  NYL.  var.  pruinosa  AUCT. 
With  last  on  same  substrata,  rare.     Mankato  (Rapidan),  June 

28,  1899,  no-  I7°-      Granite  Falls,  July  14,  1899,  no.  514. 

105.  Rinodina  oreina  (Acn.)  MASS. 

On  granitic  rocks,  quartzite  and  catlinite,  abundant.  Man- 
kato, June  30,  1899,  no.  201.  New  Ulm,  July  5,  1899,  no. 
245.  North  Redwood,  July  10,  1899,  no-  3^7-  Pipestone,. 
July  18,  1899,  nos-  6°2>  603  and  605. 

106.  Rinodina  sophodes  (Acn.)  NYL. 

On  trees,  old  wood  and  rocks,  abundant.  Mankato,  June  23 > 
1899,  no<  33'  June  24>  I&99>  no.  93,  and  June  26,  1899,  no. 
115.  New  Ulm,  July  5,  1899,  nos<  2^7»  295  anc^  29%-  Granite 
Falls,  July  u,  1899,  nos.  427  and  428,  and  July  13,  1899,  nos. 
469,  486,  487  and  491. 


320  MINNESOTA    BOTANICAL    STUDIES. 

107.  Rinodina  sophodes  (Acn.)  NYL.  var.  tephraspis  TUCK. 
On  quartzite,  rare.     Pipestone,  July  18,  1899,  no.  632. 
Not  previously  reported  from  Minnesota. 

108.  Rinodina  sophodes  (Acn.)  NYL.  var.  exigua  FR. 

On  old  wood,  locally  common.  Mankato,  June  22,  1899, 
no.  22.  Granite  Falls,  July  u,  1899,  no.  434. 

109.  Rinodina  bischoffii  (HEPP.)  KOERB. 

On  limestone  and  granite,  rare.     Mankato,  June   29,  1899, 
no.  194.     Morton,  July  7,  1899,  no.  316. 
Not  previously  reported  from  Minnesota. 

no.  Rinodina  lecanorina  MASS. 

On  boulders,  rare.     Mankato,  June  26,  1899,  no.  127. 

Not  previously  reported  from  Minnesota  and  new  to  North 
America. 

in.  Pertusaria  velata  (TURN.)  NYL. 

On  trees,  rare.  Mankato  (Minneopa  Falls),  June  26,  1899, 
no.  135. 

112.  Pertusaria  pustulata  (Acn.)  NYL. 

On  trees,  rare.  Mankato,  June  23,  1899,  no.  30,  and  July 
i,  1899,  no.  214. 

113.  Pertusaria  leioplaca  (AcH.)  SCHAER. 

On  trees,  rare.     Mankato,  June  23,  1899,  no.  68. 

114.  Urceolaria  scruposa  (L.)  NYL. 

On  earth  and  rocks,  infrequent.  Mankato,  June  26,  1899, 
no.  128.  Mankato  (Rapidan),  June  28,  1899,  no.  187.  Red- 
wood Falls,  July  8,  1899,  no.  332.  Granite  Falls,  July  n, 
1899,  no.  393.  Pipestone,  July  19,  1899,  no-  ^4°- 

115.  Urceolaria  actinostoma  PERS. 

On  granite,  rare.     Granite  Falls,  July  u,  1899,  no.  416. 
Not  previously  reported  from  Minnesota. 

116.  Stereocaulon  paschale  (L.)  FR. 

On  mossy  rocks,  only  seen  once  in  small  quantity.  Red- 
wood Falls,  July  8,  1899,  no.  359. 

117.  Cladonia  symphycarpia  FR.  var.  epiphylla  (Acn.)  NYL. 
On  earth,  rare.      Mankato,  June  26,  1899,  no.  108. 

118.  Cladonia  mitrula  TUCK. 

On  earth,  rare.  Mankato,  June  26,  1899,  no.  98.  Granite 
Falls,  July  n,  1899,  no.  436.  The  last  a  small  form  approach- 
ing Cladonia  c&sj)iticia  (Pers.)  Fl. 


Fink  :     LICHENS    OF    THE    MINNESOTA    VALLEY.  321 

119.  Cladonia  cariosa  (Acn.)  SPRENG. 

On  earth,  rare.     Mankato,  June  26,  1899,  no.  103. 
Redwood  Falls,  July  8,  1899,  no.  336.     Granite  Falls,  July  17, 
1899,  no.  587. 

120.  Cladonia  pyxidata  (L.)  FR. 

On  earth,  common  or  frequent.  Mankato,  June  26,  1899,  no. 
104.  Mankato  (Rapidan),  June  28,  1899,  no.  168.  New  Ulm, 
July  5,  1899,  nos.  272  and  276,  July  n,  1899,  no.  397,  July  12, 
1899,  no,  453,  and  July  17,  1899,  no.  562.  Pipestone,  July  18, 
1899,  no.  627. 

121.  Cladonia  fimbriata  (L.)  FR. 

On  earth,  rare.  Mankato,  June  26,  1899,  no  123.  Granite 
Falls,  July  12,  1899,  no.  582.  Pipestone,  July  18,  1899,  no. 
604. 

122.  Cladonia  fimbriata  (L.)  FR.  var.  tubaeformis  FR. 

On  old  wood  and  earth,  rare.  Mankato,  June  26,  1899,  no. 
124.  New  Ulm,  July  5,  1899,  no.  279.  Granite  Falls,  July 
n,  1899,  no.  425,  July  13,  1899,  no-  495>  and  July  i7i  l899> 
nos.  551  and  563. 

123.  Cladonia  fimbriata  (L.)  FR.  var.  radiata  FR. 

On  earth,  rare.     Redwood  Falls,  July  8,  1899,  no-  337- 

124.  Cladonia  gracilis  (L.)  NYL. 

On  old  wood  and  earth,  frequent  at  Mankato,  elsewhere  rare. 
Mankato,  June  22,  1899,  no-  4»  and  June  2^>  1899,  no.  100. 
Granite  Falls,  July  13,  1899,  nos.  468  and  488,  and  July  17, 
1899,  no.  556. 

125.  Cladonia  gracilis  (L.)  NYL.  var.  symphycarpia  TUCK. 
On  old  wood,  rare.     Mankato,  June  26,  1899,  no.  99. 

126.  Cladonia  gracilis  (L.)  NYL.  var.  verticillata  FR. 

On  earth,  rare.  Mankato,  June  26,  1899,  no>  IO1-  Granite 
Falls,  July  17,  1899,  no-  557- 

127.  Cladonia  gracilis  (L.)  NYL.  var.  hybrida  SCHAER. 

On  earth,  rare.  Mankato  (Rapidan),  June  28,  1899,  no. 
158.  Redwood  Falls,  July  8,  1899,  no.  347. 

128.  Cladonia  turgida  (EHRH.)  HOFFM. 

On  earth,  rare.     New  Ulm,  July  5,  1899,  no.  253. 

129.  Cladonia  caespiticia  (PERS.)  FL. 

On  earth,  rare.  Redwood  Falls,  July  8,  1899,  no.  342. 
Granite  Falls,  July  17,  1899,  no.  555. 


322  MINNESOTA    BOTANICAL    STUDIES. 

130.  Cladonia  furcata  (Huos.)  FR. 

On  earth,  rare.  Mankato  (Minneopa  Falls),  June  27,  1899., 
nos.  155  and  157. 

131.  Cladonia  furcata  (HUBS.)  FR.  var.  racemosa  FL. 

On  earth  in  shaded  places,  rare.  Mankato  (Minneopa  Falls),. 
June  27,  1899,  no.  156.  Redwood  Falls,  July  8,  1899,  no.  334. 

132.  Cladonia  furcata  (HUBS.)  FR.  var.  pungens  FR. 

On  earth,  rare.     Redwood  Falls,  July  6,  1899,  no.  303. 

133.  Cladonia  rangiferina  (L.)  HOFFM. 

On  earth,  frequent  locally  among  granitic  rocks.  New  Ulm, 
July  5,  1899,  no.  252. 

134.  Cladonia  rangiferina    (L.)  HOFFM.  var.  sylvatica  L. 
On  earth,  rare.     Redwood  Falls,  July  8,  1899,  no.  358. 

135.  Cladonia  macilenta  (EHRH.)  HOFFM. 

On  old  wood,  rare.     Mankato,  June  29,  1899,  no.  195. 

136.  Cladonia  cristatella  TUCK. 

On  old  stumps,  rare.  Mankato  (Rapidan),  June  28,  1899, 
no.  162.  Redwood  Falls,  July  8,  1899,  no.  356.  Granite  Falls, 
July  13,  1899,  no-  467  and  July  17,  1899,  no.  575. 

137.  Cladonia  cristatella  TUCK.  var.  paludicola  TUCK. 

Once  collected  on  an  old  log.  Mankato,  June  26,  1899,  no. 
122.  Squamules  not  powdery. 

Not  previously  reported  from  Minnesota,  and  new  to  the  upper 
Mississippi  valley. 

138.  Biatora  decipiens  (EHRH.)  FR. 

Common    on    earth    containing    calcareous     drift    pebbles. 
Granite  Falls,  July  13,  1899,  no.  500. 
Not  previously  reported  from  Minnesota. 

139.  Biatora  decipiens  (EHRH.)  FR.  var.  dealbata  AUCT. 
Common   on   earth   with  the   last.     Granite   Falls,  July   13, 

1899,  no.  499. 

Not  previously  reported  from  Minnesota. 

140.  Biatora  icterica  MONT. 

On  earth,  rare.  Granite  Falls,  July  n,  1899,  no.  398,  and 
July  18,  1899,  no.  519. 

141.  Biatora  rufonigra  TUCK. 

On  granitic  rocks  and  quartzite,  common.  New  Ulm,  July  5, 
1899,  no-  265-  Morton,  July  7,  1899,  no-  325-  Granite  Falls, 
July  12,  1899,  no.  454. 


Fink  :     LICHENS    OF    THE    MINNESOTA    VALLEY.  323 

142.  Biatora  coarctata  (SM.,  NYL.)  TUCK. 

On  limestone  and  sandstone,  rare.  Mankato,  June  26,  1899, 
no.  113.  Mankato  (Rapidan),  June  28,  1899,  no.  173. 

1420;.  Biatora   coarctata  (SM.,  NYL.)   TUCK.  var.  brugeriana, 

SCHAER. 

On  sandstone,  locally  abundant.  Mankato  (Minneopa  Falls), 
June  27,  1899,  nos.  139,  142,  145  and  146.  Mankato  (Rapi- 
dan), June  29,  1899,  nos.  172,  174  and  176. 

143.  Biatora  uliginosa  (SCHRAD.)  FR. 

On  earth,  infrequent.  Mankato,  June  26,  1899,  no.  128. 
New  Ulm,  July  5,  1899,  no.  250. 

144.  Biatora  myriocarpoides  (NYL.)  TUCK. 

On  quartzite,  locally  common.  New  Ulm,  July  5,  1899,  no. 
300. 

145.  Biatora  varians  (Acn.)  TUCK. 

On  trees,  probably  frequent  locally.  Granite  Falls,  July  15, 
1899,  no-  502- 

146.  Biatora  flexuosa  FR. 

On  dead  cedar,  rare.     Granite  Falls,  July  13,  1899,  no.  477. 
Not  previously  reported  from  Minnesota. 

147.  Biatora  hypnophila  (TURN.)  TUCK. 

On  earth  and  limestone,  rare.  Mankato,  June  23,  1899,  no. 
36,  and  June  26,  1899,  no.  120. 

148.  Biatora  naegelii  HEPP. 

On  trees,  infrequent.  Granite  Falls,  July  13,  1899,  no. 
484,  and  July  15,  1899,  no-  53°- 

149.  Biatora  rubella  (EHRH.)  RABENH. 

On  trees,  common  locally.  Mankato,  June  23,  1899,  no.  35, 
and  June  26,  1899,  no.  130.  Mankato  (Minneopa  Falls),  June 
27,  1899,  no.  138. 

150.  Biatora  fuscorubella  (HOFFM.)  TUCK. 

On  trees  and  rocks,  common  at  Mankato,  else  where  rare  or 
infrequent.  Mankato,  June  23,  1899,  no-  29-  Mankato  (Min- 
neopa Falls),  June  27,  1899,  no.  141.  New  Ulm,  July  5,  1899, 
no.  274.  Granite  Falls,  July  15,  1899,  nos.  529  and  536. 

151.  Biatora  suffusa  FR. 

On  trees,  rare.  Mankato,  June  23,  1899,  no.  37.  Granite 
Falls,  July  15,  1899,  no.  540. 

Not  previously  reported  from  Minnesota. 


324  MINNESOTA    BOTANICAL    STUDIES. 

152.  Biatora  muscorum  (Sw.)  TUCK. 

On  earth,  frequent  and  once  on  sandstone.  Mankato,  July 
i,  1899,  no-  22°-  New  Ulm,  July  4,  1899,  no-  237-  Red- 
wood Falls,  July  8,  1899,  no.  344.  Granite  Falls,  July  n, 
1899,  no.  433.  Pipestone,  July  18,  1899,  nos.  590  and  591. 

153.  Biatora  inundata  FR. 

On  limestone  and  sandstone,  common.  Mankato,  June  22, 
1899,  no.  24.  New  Ulm,  July  4,  1899,  no.  234. 

154.  Lecidea  enteroleuca  FR. 

On  trees,  common  at  Granite  Falls.  Mankato,  June  23,  1899, 
no.  80,  and  July  i,  1899,  no-  2O9-  New  Ulm,  July  4,  1899,  no- 
231.  Redwood  Falls,  July  8,  1899,  no.  354.  Granite  Falls, 
July  n,  1899,  no-  429>  JulY  Z3»  l899»  nos.  476,  479,  485  and 
496,  and  July  14,  1899,  no.  516. 

155.  Lecidea  enteroleuca  FR.  var.  achrista  SOMMERF. 

On  trees,  frequent.  Granite  Falls,  July  13,  1899,  nos.  471 
and  475,  and  July  15,  1899,  no-  546- 

156.  Buellia  spuria  (SCHAER.)  ARN. 

On  granitic  rocks,  quartzite  and  pipestone,  frequent  or  com- 
mon. New  Ulm,  July  5,  1899,  no.  294.  Morton,  July  7,  1899, 
no.  324.  Granite  Falls,  July  12,  1899,  nos.  445  and  458. 
Pipestone,  July  18,  1899,  no.  612. 

157.  Buellia  alboatra  (HOFFM.)  TH.  FR. 

On  trees,  especially  Ulmus,  rare  at  first  locality  and  more 
common  at  second.  Mankato,  June  22,  1899,  no.  15.  Gran- 
ite Falls,  July  n,  1899,  no.  382,  and  July  17,  1899,  no.  577. 

158.  Buellia  alboatra  (HOFFM.)  TH.  FR.  var.  saxicola  FR. 

On  limestone,  shaded,  rare.     Mankato,  June  22,  1899,  no.  16. 
Not  previously  reported  from  Minnesota. 

159.  Buellia  parasema  (Acn.)  TH.  FR. 

On  trees,  infrequent.  Mankato,  June  23,  1899,  no.  34, 
and  June  24,  1899,  no.  85.  New  Ulm,  July  3,  1899,  no<  224' 
Granite  Falls,  July  n,  1899,  no.  388. 

160.  Buellia  myriocarpa  (DC.)  MUDD. 

On  old  wood,  common  or  frequent.  Mankato,  June  22, 
1899,  no.  20.  Mankato  (Rapidan),  June  28,  1899,  no.  184. 
Granite  Falls,  July  17,  1899,  no.  552. 


Fink  :   LICHENS  OF  THE  MINNESOTA  VALLEY.  325 

161.  Buellia  pullata  TUCK. 

On  rocks,  frequent.  Morton,  July  7,  1899,  no.  327.  North 
Redwood,  July  10,  1899,  nos'  3^5  anc^  3^-  Granite  Falls, 
July  n,  1899,  no.  405.  Pipestone,  July  18,  1899,  no.  600. 

Not  previously  reported  from  Minnesota. 

162.  Buellia  turgescens  (NYL.)  TUCK. 

On  old  boards,  rare.     Mankato,  June  22,  1899,  no.  21. 
Not  previously  reported  from  Minnesota. 

163.  Buellia  petraea  (FLOT.,  KOERB.)  TUCK. 

On  granite,  quartzite  and  pipestone,  abundant.  New  Ulm, 
July  5,  1899,  nos.  242  and  293.  Redwood  Falls,  July  6,  1899, 
no.  306.  Granite  Falls,  July  11,  1899,  no>  422> 

164.  Buellia  petrasa  (FLOT.,  KOERB.)  TUCK.   var.    montagnaei 
TUCK. 

On  same  rocks  as  last  and  even  more  abundant ;  however, 
only  a  single  collection  on  a  bowlder  at  first  locality.  Man- 
kato (Rapidan),  June  28,  1899,  no.  183.  Morton,  July  7,  1899, 
nos.  318  and  321.  North  Redwood,  July  10,  1899,  no.  362. 
Granite  Falls,  July  n,  1899,  no.  4223.  Pipestone,  July  18, 
1899,  nos.  593  and  620. 

165.  Opegrapha  varia  (PERS.)  FR. 

On  trees,  abundant.  Mankato,  June  23,  1899,  no.  40,  June 
26,  1899,  no.  no,  and  July  i,  1899,  no.  204.  Granite  Falls, 
July  n,  1899,  nos.  419  and  421,  Julv  13,  1899,  no<  4^3>  an^ 
July  15,  1899,  nos.  539  and  548. 

166.  Opegrapha  varia  (PERS.)  FR.  var.  pulicaris  (HOFFM.)  FR. 
On  trees,  rare.     Granite  Falls,  July  15,  1899,  no-  52^- 

Not  previously  reported  from  Minnesota. 

167.  Graphis  scripta  (L.)  ACH. 

On  trees,  common  at  Mankato.  Mankato,  June  22,  1899,  no. 
14.  Granite  Falls,  July  13,  1899,  no.  503. 

168.  Graphis  scripta  (L.)  ACH.  var.  recta  (HUMB.)  NYL. 

On  birches,  rare.  Mankato  (Minneopa  Falls),  June  27,  1899, 
no.  1413. 

169.  Graphis  scripta  (L.)  ACH.  var.  limitata  ACH. 

On  trees,  rare.  Mankato,  June  22,  1899,  no.  23.  Granite 
Falls,  July  13,  1899,  no  469,  and  July  15,  1899,  no.  538. 

170.  Arthonia  lecideella  NYL. 

On  trees,  infrequent.     Mankato,  June  23,  1899,  no.  69. 


326  MINNESOTA    BOTANICAL    STUDIES. 

/ 

171.  Arthonia  dispersa  (SCHRAD.)  NYL. 

On  trees,  common.  Mankato,  June  22,  1899,  no.  25,  and 
July  23,  1899,  no.  72.  Granite  Falls,  July  13,  1899,  no.  490. 

172.  Arthonia  radiata  (PERS.)  TH.  FR. 

On  trees,  infrequent.  Mankato,  June  23,  1899,  no.  70. 
Granite  Falls,  July  13,  1899,  no.  489,  and  July  15,  1899,  no. 
532- 

173.  Arthonia  ptmctiformis  ACH. 

On  maples,  rare.      Granite  Falls,  July  15,  1899,  no.  341  a. 

i73<z.  Arthonia  sp. 

On  trees,  rare.     Granite  Falls,  July  15,  1899,  no.  541. 
With    general   appearance   of  Arthonia    dispersa    (Schrad.) 

22 26 

Nyl.,  but  the  colorless  spores  are  four  celled  and  -  -  mic. 

7-5~8-5 
Not  previously  reported  from  Minnesota. 

174.  Calicium  parietinum  ACH. 

On  old  wood,  probably  rare.  Mankato,  June  22,  1899,  no. 
8,  and  June  24,  1899,  no.  87,  Redwood  Falls,  July  8,  1899,  no. 
326. 

175.  Calicium  quercinum  PERS. 

Collected  once  only,  on  cedar.  Granite  Falls,  July  13,  1899, 
no.  478. 

176.  Coniocybe  pallida  (PERS.)  FR. 

On  a  large  oak,  only  once  collected.  Mankato,  July  7,  1899, 
no.  206. 

177.  Endocarpon  miniatum  (L.)  SCHAER. 

Abundant  on  limestone  bluffs,  frequent  on  granite  and  rare  on 
quartzite.  Mankato,  June  23,  1899  no.  46  and  June  25,  1899, 
no.  59.  Redwood  Falls,  July  6,  1899,  no.  312.  Granite  Falls, 
July  n,  1899,  no.  373,  and  July  13,  1899,  no.  474.  Pipestone, 
July  18,  1899,  no-  606. 

178.  Endocarpon   miniatum   (L.)   SCHAER.    var.    complicatum 
SCHAER. 

On  substrata  noted  above  and  also  on  pipestone,  frequent. 
Mankato,  June  25,  1899,  no.  57.  New  Ulm,  July  5,  1899,  no. 
249.  North  Redwood,  July  10,  1899,  no.  363.  Granite  Falls, 
July  12,  1899,  no.  448,  and  July  13,  1899,  no.  495.  Pipestone, 
July  18,  1899,  no.  615,  and  July  19,  1899,  no>  ^39- 


Fink :   LICHENS  OF  THE  MINNESOTA  VALLEY.  327 

179.  Endocarpon  fluviatile  DC. 

On  rocks  frequently  wet,  infrequent.  Morton,  July  7,  1899, 
no.  322.  Granite  Falls,  July  12,  1899,  4483. 

180.  Endocarpon  arboreum  SCHWEIN. 

On  trees  and  shaded  rocks,  once  seen  on   each.     Redwood 
Falls,  July  6,  1899,  no.  308,  and  July  8,  1899,  no>  339- 
Not  previously  reported  from  Minnesota. 

181.  Endocarpon  hepaticum  ACH. 

On  earth  and  sandstone,  common.  Mankato  (Rapidan), 
June  28,  1899,  no-  Z75-  New  Ulm,  July  4,  1899,  no>  235>  ant* 
July  5,  1899,  no.  244.  North  Redwood,  July  10,  1899,  no. 
364.  Granite  Falls,  July  n,  1899,  no-  3^4-  Pipestone,  July 
18,  1899,  no-  623. 

182.  Endocarpon  pusillum  HEDW. 

On  limestone  bluffs,  sandstone,  calcareous  drift  pebbles,  and 
once  on  earth,  common.  Mankato,  June  23,  1899,  no.  39. 
New  Ulm,  July  4,  1899,  no.  236.  Granite  Falls,  July  12, 

1899,   no-  442»  JulY  T3»   l899>  no-  5°9»   and  July   J4»   l899» 
no.  526. 

183.  Endocarpon  pusillum  HEDW.  var.  garovaglii  KPH. 

On  earth  and  sandstone,  frequent.  Mankato,  July  i,  1899, 
no.  219.  Mankato  (Rapidan),  June  28,  1899,  no.  186.  New 
Ulm,  July  5,  1899,  no.  282.  Morton,  July  7,  1899,  no.  317. 
Pipestone,  July  18,  1899,  no.  616. 

184.  Thelocarpon  prasinellum  NYL. 

On  old  wood  and  sandstone,  frequent.  Mankato,  June  22, 
1899,  no-  6»  and  June  26,  1899,  no.  132. 

I  cannot  bring  that  on  sandstone  under  any  of  the  rock  species, 
and  it  seems  to  belong  here. 

185.  Staurothele  umbrina  (WAHL.)  TUCK. 

On  granite,  limestone  and  quartzite,  frequent.  Mankato, 
June  23,  1899,  no-  82.  Granite  Falls,  July  u,  1899,  no.  412, 
July  15,  1899,  no-  547'  and  July  17,  1899,  no.  566. 

186.  Staurothele  diffractella  (NYL.)  TUCK. 

On  sandstone,  granite,  quartzite  and  calcareous  drift  pebbles, 
rare.  New  Ulm,  July  4,  1899,  nos.  238  and  240.  Granite 
Falls,  July  13,  1899,  no.  501,  and  July  17,  1899,  no.  560. 

Not  previously  reported  from  Minnesota. 


328  Fink :  LICHENS  OF  THE  MINNESOTA  VALLEY. 

187.  Staurothele  drummondii  TUCK. 

On  granite,  quartzite  and  pipestone,  common  in  damp  places 
at  Granite  Falls  and  Pipestone.  Redwood  Falls,  July  8,  1899, 
no.  327.  Granite  Falls,  July  12,  1899,  no.  457,  July,  13,  1899, 
no.  494,  and  July  17,  1899,  no.  554.  Pipestone,  July  18,  1899, 
nos.  595,  619,  622  and  628. 

"188.  Verrucaria  fuscella  FR. 

On  limestone,  infrequent.  Mankato,  June  23,  1899,  no-  42' 
i88«.  Verrucaria  nigrescens  PERS. 

On  limestone  common  and  once  seen  on  a  granite  bowlder. 
Mankato,  June  23,  1899,  no.  65,  June  24,  1899,  no.  96,  and 
June  26,  1899,  no.  116. 

189.  Verrucaria  muralis  ACH. 

On  limestone  in  bluffs  and  drift  pebbles,  abundant  at  Man- 
kato. Mankato,  June  22,  1899,  no.  26.  Granite  Falls,  July 
13,  1899,  no.  511,  and  July  14,  1899,  no.  525. 

190.  Pyrenula  punctiformis  (ACH.)  NAEG. 

On  trees,  infrequent.     Mankato,  June  24,  1899,  nos.  90  and  94. 

191.  Pyrenula  punctiformis  (AcH.)  NAEG.  var.  fallax  NYL. 
On  birch,  infrequent.     Mankato,  June   24,  1889,  no.  66,  and 

June  26,  1899,  no.  109. 

192.  Pyrenula  gemmata  (AcH.)  NAEG. 

On  trees  frequent.     Granite  Falls,  July  14,  1899,  no'  5I3< 
Not  previously  reported  from  Minnesota. 

193.  Pyrenula  hyalospora  NYL. 

On  trees,  probably  rare.     Mankato,  June   23,  1899,  no.  32, 
and  June   25,  1899.     Granite  Falls,  July  13,  1899,  no.  470. 
Not  previously  reported  from  Minnesota. 

194.  Pyrenula  nitida  ACH. 

On  trees,  rare.  Mankato,  July  i,  1899,  no.  211.  New  Ulm, 
July  3,  1899,  no.  222. 

195.  Pyrenula  thelena  (Acn.)  TUCK. 

On  birch,  common.  Mankato  (Minneopa  Falls),  June  27, 
1899,  no.  140. 

196.  Pyrenula  cinerella  (FLOT.)  TUCK. 

On  trees,  infrequent.  Mankato  (Minneopa  Falls),  June  27, 
1899,  no.  143. 

Spores  reaching  12-16  by  6-8  mic.  in  one  collection.  Thus 
larger  than  usual  American  forms. 


Fink  :   LICHENS  OF  THE  MINNESOTA  VALLEY.  329 

197.  Pyrenula  cinerella  (PLOT.)  TUCK.  var.   quadriloculata  var. 
nov. 

On  birch,  probably  rare.  Mankato,  June  26,  1899,  no.  129. 
Mankato  (Rapidan),  June  28,  1899,  no.  163%. 

Second  time  collected  in  Minnesota  and  both  times  from 
hosts  of  same  genus. 

198.  Pyrenula  quinqueseptata  (NYL.)  TUCK. 

On  trees,  rare.     Mankato,  July  i,  1899,  no>  2O^' 
Spores  frequently  showing  8  cells,  which  is  not  common  for 
the  species. 

Not  previously  reported  from  Minnesota. 

199.  Pyrenula  leucoplaca  (WALLR.)  KBR. 

On  trees,  common.  Mankato,  June  23,  1899,  no.  31.  Man- 
kato (Rapidan),  June  28,  1899,  no-  I^°-  Mankato,  July  i, 
1899,  no.  207.  Granite  Falls,  July  13,  1899,  no>  4^4»  and  July 
15,  1899,  nos.  527,  535  and  543. 

200.  Pyrenula  glabrata  (AcH.)  MASS. 

On  trees,  rare.     Mankato,  June  24,  1899,  no.  88. 
Not  previously  reported  from  Minnesota. 

201.  Pyrenula  megalospora  sp.  nov. 

Thallus  rather  smooth,  indeterminate,  prominent,  gray  or 
grayish  white.  Apothecia  scattered  or  occasionally  aggregated 
in  clusters  of  two  or  three,  black  or  brownish  black,  convex 
with  the  ostiole-bearing  apex  somewhat  pointed,  semi-immersed 
or  becoming  more  superficial,  .4  to  .75  mm.  in  diameter.  Am- 
phithecium  white.  Paraphyses  capillary  and  very  distinct. 
Asci  cylindrical,  .25  to  .3  mm.  in  length.  Spores  colorless,  2- 
celled,  oblong  with  ends  obtuse  or  somewhat  pointed,  somewhat 
constricted  at  the  septum,  large  for  2-celled  spores  of  the  genus 
(35-60  by  14-21  mic.),  8  in  asci,  crowded  and  obliquely  uni- 
seriate. 

On  trees,  frequent.  Mankato,  June  26,  1899,  no.  112  and 
July  i,  1899,  no.  209.  Granite  Falls,  July  n,  1899,  no.  381 
and  July  n,  1899,  no.  576. 


XX.     A    SYNONYMIC     CONSPECTUS     OF    THE 

NATIVE   AND    GARDEN   AQUILEGIAS    OF 

NORTH    AMERICA. 


K.  C.  DAVIS. 


The  name  Aquilegia  (Linn.  Sp.  PL  533,  1753)  is  probably 
not  from  aquila,  eagle,  as  commonly  given,  but  from  aquilegus, 
water-drawer.  The  name  may  have  been  applied  from  the 
supposed  power  of  the  roots  to  extract  water  from  rocks,  among 
which  they  so  often  grow.  They  are  commonly  called  Colum- 
bines. 

Hardy  perennial  herbs,  mostly  with  paniculate  branches  ter- 
minated by  showy  flowers  ;  leaves  1-3  times  ternately  compound, 
commonly  glaucous ;  leaflets  roundish  and  obtusely  lobed : 
flowers  large,  showy,  appearing  usually  in  spring  or  early 
summer;  sepals  5,  regular,  petaloid  ;  petals  concave,  produced 
backward  between  the  sepals  forming  a  hollow  spur ;  stamens 
numerous  ;  fruit  of  about  5  many-seeded  follicles. 

About  30  species  are  distinct ;  all  of  the  north  temperate  re- 
gions of  the  world.  Nearly  half  of  these  (12)  are  natives  of 
North  America.  Most  of  the  native  species  and  varieties  are 
used  in  American  and  European  gardens,  and  ten  foreign  spe- 
cies have  already  been  introduced  here.  Aquilegia  furnishes 
many  useful,  ornamental  forms  eminently  fitted  for  choice  mixed 
borders  and  beds.  A  good,  deep,  rather  sandy,  well  drained 
soil  is  the  best.  Seeds  sown  in  pans,  in  cold  frames  in  March, 
or  open  air  in  April,  occasionally  bloom  the  first  season,  but 
generally  the  second.  The  seed  germinates  slowly,  and  the 
ground  should  be  kept  moist  on  top  during  this  period.  The 
different  species  should,  if  possible,  be  kept  some  distance  apart 
if  pure  seed  is  desired,  as  the  most  divers  species  hybridize 
directly.  They  may  be  propagated  by  root  division  but  better 
by  seed.  Absolutely  pure  seed  is  hard  to  obtain  except  from 
the  plants  in  the  wild  state ;  and  some  of  the  mixed  forms  are 


332  MINNESOTA    BOTANICAL    STUDIES. 

quite  inferior  to  the  true  species  from  which  they  have  come. 
A.  ccerulea,  A.  glandulosa,  and  A.  vulgar  is  are  apt  to  flower 
only  two  or  three  years  from  the  same  plant  and  should  be 
treated  as  biennials ;  but  A.  vulgaris  may  be  kept  active  for  a 
longer  period  by  transplanting. 

The  latest  extended  accounts  of  species  in  this  group  are  by 
J.  G.  Baker,  in  Gardener's  Chronicle,  1878  ;  and  B.  L.  Robin- 
son, in  Gray's  Synoptical  Flora,  i  :  42-45,  1895. 

In  presenting  this  and  the  accompanying  paper,  I  desire  to 
extend  thanks  to  all  who  have  so  kindly  aided  me  in  my  studies 
of  Ranunculaceous  genera ;  particularly  I  am  under  obligations 
to  Professor  L.  H.  Bailey  for  many  valuable  suggestions  at 
times  of  greatest  need  and  for  placing  about  me  the  largest  col- 
lection of  colored  plant  portraits  and  the  largest  garden  her- 
barium in  America,  and  to  Professor  W.  W.  Rowlee  for  placing 
at  my  disposal  not  only  the  entire  collection  of  herbarium  speci- 
mens in  the  department  of  botany  of  Cornell  University,  but  also 
numerous  living  roots  and  plants  from  which  to  better  study 
vegetative  characters. 

KEY  TO  SPECIES  OF  AQUILEGIA. 

A.   Sepals  not  more  than  ^  or  ^  inch  long;  expanded  flower  i  or 

\y>z  inches  in  diameter. 
B.   Limb  of  petal  shorter  than  the  sepal. 

C.   Stem-leaves  present;   stem  ij^-to  2^  feet  high. 

D.   Spur  straight,  not  knobbed lactiflora. 

DD.   Spur  knobbed,  bent  inward oxysepala. 

CC.  Stem-leaves  wanting ;   stem  reduced  to  a  short  scape. 

Jonesii. 
BB.   Limb  of  petal  about  equal  to  the  sepal. 

C.  Leafless  or  nearly  so ;   stem  scapiform elegantula. 

CC.   Leaves  two  or  more  on  a  stem. 

D.   Plant  low,   slender   (commonly  6-S  inches)  ;   spurs 

incurved. 
E.   Leaves,  stems  and  follicles  pubescent. 

brevistyla. 
EE.   Leaves,  stems  and  follicles  smooth. 

saximontana. 

DD.   Plant  one  foot  high  or  more ;  spurs  nearly  straight. 
E.   Stamens  protruding  beyond  the  petal-limbs. 

F.   Spurs  somewhat  knobbed Canadensis. 

FF.   Spurs  not  knobbed viridiflora. 


Davis:   A  SYNONYMIC  CONSPECTUS  OF  AOJJILEGIAS.         333 

EE.   Stamens   hardly  protruding   or    shorter   than 

petal-limb. 
F.  Plant  finely  pubescent  above. 

Buergeriana. 
FF.  Plant     glandular -pubescent    and    viscid 

above mtcrantha. 

AA.   Sepals  about  one  inch  long;  expanded  flower  about  two  inches 

in  diameter. 

B.   Spurs  shorter  than  the  petal  limb,  and  incurved — Jlabellata. 
BB.   Spurs  at  least  as  long  as  the  petal-limb. 
C.   Stamens  short,  not  much  protruding. 

D.   Spurs  only  slightly  curved,  not  knobbed... leptoceras. 
DD.   Spurs  much  incurved  or  coiled. 

E.   Follicles  densely  pubescent vulgaris. 

EE.  Follicles  glabrous Sibirica. 

CC.   Stamens  long,  protruding  far  beyond  the  petal-limb. 
D.   Sepals  green,  keeled  ;  young  fruits  winged. 

Skinneri. 
DD.   Sepals  usually  yellow  or  red,  not  keeled;   fruits 

never  winged formosa. 

BBB.   Spurs  very  long,  often  several  times  as  long  as  petal-limbs. 

C.   Length  of  spur  about  i  or  i^  inches pubescens. 

CC.   Length  of  spur  about  2  inches chrysantha. 

CCC.   Length  of  spur  4  inches  or  more longissima. 

AAA.   Sepals   i%   to  i^    or  even  2  inches  long;   expanded  flower 
2  ^2  to  3  inches  in  diameter;   stamens  not  protruding. 

B.   Spurs  long  and  not  incurved cczrulea. 

BB.   Spurs  incurved  and  not  longer  than  the  petal-limbs. 

C.   Follicles  few,  pubescent alptna. 

CC.   Follicles  6-10;  densely  hairy .glandulosa. 

A.  lactiflora  KAR.  &  KIR.  in  Mosc.  Bull.  15:  374.     1841. 

Stem  i  y2  feet  high,  glabrous  in  the  lower  part :  partial-peti- 
oles of  root  leaves  i  ^  to  2  inches  long,  leaflets  sessile  or  short 
stalked,  i  inch  long,  many  lobes  reaching  half  way  down ; 
stem  leaves  petioled  and  compound  ;  flowers  about  3  on  a  stem  ; 
sepals  nearly  white,  or  tinged  with  blue,  over  y2  inch  long, 
narrow ;  petal-limb  half  as  long  as  the  sepal ;  spur  ^  inch, 
slender,  nearly  straight,  not  knobbed  at  tip ;  stamens  equal  in 
length  to  the  limb.  June.  Altai  Mts.,  Siberia.  A  desirable 
species  but  not  much  used  in  gardens. 

A.  oxysepala  TRAUT.  &  MEYER,  in  Middend.  Reise.  Florula 
Ocho.  Phaen.  10.  1856. 


334  MINNESOTA    BOTANICAL    STUDIES. 

Plant  2^  feet  high,  slightly  pubescent  above  :  radical  leaves 
long-petioled,  secondary  divisions  sessile  :  sepals  ovate-lanceo- 
late, much  exceeding  the  petal-limbs  in  length,  which  are  6 
lines  long,  white,  rounded-truncate ;  stamens  not  protruding 
beyond  the  petal-limb  ;  spur  knobbed,  bent  inward,  shorter  than 
petal-limb  :  follicles  pubescent  with  styles  their  own  length. 
June.  Eastern  Siberia.  In  1898  F.  H.  Horsford  said  of  this  : 
"  the  first  to  bloom  with  me,  and  one  of  the  most  attractive  in 
the  list.  It  is  one  of  the  most  dwarfed  ;  flowers  large,  blue, 
yellow  and  white  :  it  comes  so  much  before  the  others  that  its 
capsules,  as  a  rule,  all  fertilize  before  any  of  the  other  species 
come  into  flower." 

A.  Jonesii  PERRY,  Am.  Nat.  8  :   211.     1874. 

True  stem  very  short  or  almost  wanting,  soft-pubescent : 
tufted  root-leaves  an  inch  or  two  high  from  the  stout,  ascending 
branches  of  the  rootstock ;  biternately  divided  partial  petioles 
very  short  or  none  ;  leaflets  very  crowded  :  flowers  blue  ;  sepals 
oblong,  obtuse,  equalling  the  spurs  and  twice  the  length  of  the 
petal  limbs  and  head  of  stamens :  follicles  glabrous,  large, 
nearly  i  inch  long,  styles  ^  as  long ;  peduncles  lengthening  to 
about  3  inches  in  fruit.  July.  N.  W.  Wyoming,  Mont.,  to 
Brit.  Am.  Garden  &  Forest,  9:  365. 

A.  elegantula  GREENE,  Pitt.  4  :   14.     1899. 

Erect,  slender,  mostly  less  than  a  foot  high,  glabrous  except 
on  the  inflorescence,  the  peduncle  and  exterior  of  the  flowers 
hirtellous-pubescent ;  stem  scapiform,  usually  only  bracted ; 
radical  leaves  long-petioled,  glabrous  beneath :  flowers  mostly 
solitary,  terminal,  small,  about  i  inch  long,  sepals  light  green, 
erect ;  petal-limbs  light  yellow,  erect ;  spurs  straight,  longer 
than  the  sepals,  rather  widely  inflated  above  and  light  scarlet 
in  that  part;  filaments  short;  styles  exserted.  June.  Slide 
Rock  Canon,  and  in  Spruce  woods  Mt.  Hesperus,  S.  Colo. 
Described  from  the  original.  Type  in  Greene's  Herbarium  (|). 

A.  brevistyla  HOOK.  Fl.  Bor  Am.  i :  24.     1833. 

A.  vulgaris  RICHARDS,  in  Frankl.  Jl.  App.  740.     1823, 

not  Linn. 
A.  vulgaris  var.  brevistyla  GRAY,  Am.  Journ.  Sci.  Ser.  2, 

33:   242.      1862. 
A.  Laramiensis  A.  NELSON  Wyom.   Exp.   Sta.  Bull.  28: 

78.     1895-6. 


Davis:   A  SYNONYMIC  CONSPECTUS  OF  AOJJILEGIAS.         335 

Slender,  6-15  inches  high,  glandular-pubescent  above  at 
least,  several  flowered :  root-leaves  biternate,  long-petioled, 
leaflets  lobed  and  crenate ;  stem-leaves  few,  lower  ones  short- 
petioled :  flowers  blue  with  yellowish  petals,  small,  about  as 
broad  as  long  ;  petals  and  sepals  about  equal  in  length,  stamens 
a  little  shorter,  spurs  even  shorter  and  incurved :  follicles  pu- 
bescent, equalling  the  flower  in  length,  ^  inch,  becoming 
erect.  May-June.  Mts.  of  Northwest  Territory  into  South 
Dakota  (f). 

A.  saximontana  P.  A.  RYDBERG  ex  Robinson,  Syn.  Fl.  i  : 
I :  43.  1895. 

Stem  from  a  scaly  rootstock,  less  than  six  inches  high : 
leaves,  stems  and  follicles  smooth  throughout,  otherwise  like 
above.  Mts.  Colo.  (f). 

A.  Canadensis  LINN.  Sp.  PI.  533.     1753. 
A.  variegata  MOENCH.  Meth.  311.     1794. 
A.  elegans  SALISB.  Prod.  374.     1796. 

Height  i—2  feet ;  primary  divisions  of  petioles  of  root-leaves 
1-2  inches,  having  3  divisions  ;  2  or  3  of  the  stem  leaves  petioled, 
biternate  :  flowers  several  on  a  stem  ;  sepals  yellowish  or  tinted 
on  the  back  with  red,  about  ^  inch  long,  not  reflexing ;  limb 
of  petals  a  little  shorter,  yellowish,  truncate  ;  spur  ^  inch  long, 
nearly  straight,  knobbed  at  the  end,  bright  red  throughout ; 
stamens  much  protruding :  follicles  ^  inch  long,  with  style 
y^  as  long.  May— July.  Stony  banks,  etc.  East  of  the  Rocky 
Mts.  Introduced  1890.  Bot.  Mag.  246.  Loddiges'  Bot.  Cab. 
888.  Meehan's  Mo.  5  :  21. 

Var.  flaviflora  BRITTON  Bull.  Torr.  Club,  15:  97.     1888. 
.  A.flaviflora  TENNEY  Am.  Nat.  I  :  388.     1867. 
A.  c&rulea  var.  flavescens  LAWSON  Rev.   Canad.  Ranun. 

75-     1870. 

A.  flavescens  WATS.  Bot.  King.  Exp.  10.     1871. 
Flowers  a  clear  yellow.     Very  pretty.      Introduced    1889. 
Bot.  Mag.  6552  B.  (as  A.formosa  var.  flavescens]. 
Var.  depauperata  n.  var. 

A.  depauperata  JONES  Contr.  West.  Bot.  No.  8,  i.      1898. 

Stems  slender ;    leaves  and  leaflets  smaller  than  the  type ; 

flowers  also  small,  cream  colored  tinged  with  blue  and  green. 

June.     Along  streams,  and  in  Provo  Canon,  Utah.     Collected 

by  M.  E.  Jones,  who  has  the  type  (f). 


336  MINNESOTA    BOTANICAL    STUDIES. 

Var.  nana  HORT.  Plant  i  foot  high  or  less  ;  flowers  like  the 
type. 

A.  viridiflora  PALLAS  in  Act.  Petrop.  260,  t.  2.     1779. 

Stem  i-i}4  feet  high,  finely  pubescent  throughout,  several 
flowered  :  the  partial  petioles  of  root-leaves  1-2  inches  long ; 
leaflets  sessile  or  the  end  one  shortly  stalked,  lobes  rather  nar- 
row and  deep ;  lower  stem-leaves  petioled,  biternate  :  sepals 
oblong,  obtuse,  ascending,  greenish,  equalling  the  broad,  green- 
ish petal-limb,  but  not  reaching  the  head  of  stamens ;  spur 
straight,  slender,  y2  inch  long,  not  knobbed  ;  pubescent  follicles 
as  short  as  their  styles.  Summer.  East  Siberia.  Not  so  much 
used  as  the  following  variety. 

Var.  atropurpurea  n.  var. 

A.  atropurpurea  WILLD.  Enum.  Hort.  Berol.  577.      1813. 
A.  dahurica  PATR.  in  Deless.  Ic.  Sel.  t.  49.     1820. 

Limbs  of  the  petals  deep  blue  or  lilac-purple,  and  the  sepals 
and  spur  somewhat  tinged  with  the  same  hue.  Bot.  Reg.  922. 

A.  Buergeriana  SIEB.  &  Zucc.  in  Abh.  Akad.  Munch.  4 :  II, 
183.  1846. 

A.  atropurpurea  MIQUEL.  Ann.   Mus.   Lugd.   Bot.  3:  8. 

1867. 

One  foot  high,  finely  pubescent  towards  the  top  ;  branched 
to  form  several  heads,  bearing  2-3  petioled,  biternate  leaves ; 
partial-petioles  of  basal  leaves  ^-i  inch  long,  3  sessile  divi- 
sions ;  flowers  yellow  tinted  with  purple,  i-i  y2  inch  in  diam- 
eter ;  sepals  ^  inch  long,  acute,  spreading  ;  spurs  erect,  nearly 
straight,  as  long  as  the  limb  of  petal,  and  about  equalling  the 
sepal ;  head  of  stamens  equal  to  limb  in  length  :  follicles  pubes- 
cent, ^  inch  long,  style  half  as  long.  Early.  Japan. 
Brought  from  St.  Petersburg,  1892. 

A.  micrantha  EASTWOOD,  Proc.  Cal.    Acad.   Sci.   II,  4: 

559.  /.  i9.     1894. 

Stem  slender,  densely  glandular-pubescent  and  viscid  on  up- 
per parts :  leaflets  small,  cuneate,  3-cleft,  with  2-3-lobed  seg- 
ments ;  partial-petioles  of  the  lateral  leaflets  short :  flowers 
hardly  i  inch  across,  yellowish  white ;  sepals  nearly  ^  inch 
long,  nearly  half  as  broad :  petals  rather  truncate,  spur  short, 
straight,  or  slightly  curved.  July.  Canons  of  southeast  Utah 
to  Colorado  (f). 

Var.  ecalcarata  n.  var. 


Davis;   A  SYNONYMIC  CONSPECTUS  OF  AOJJILEGIAS.         337 

A.  ecalcarata  EASTWOOD  in  Zoe,    2:   226.      1891;  4:   3. 

1893. 

Flowers  sometimes  reddish,  fragrant :  spurs  sometimes  re- 
duced to  mere  sacs,  but  with  intermediated  grades  to  the  type 
form  (f). 

A.  flabellata  SIEB.  &  Zucc.  in  Abh.  Akad.  Munch.  4:  II, 
183.  1846. 

A.  vtilgaris  Thunb.  Fl.  Jap.  232.      1784.     Not  Linn. 

A.  glandulosa  Miq.  in  Ann.  Mus.  Lugd.  Bat.  3 :  8.     1867. 

Stem   i   to    i  y2    feet  high,  few  flowered :  partial-petioles  of 

root-leaves  i  inch  or  more,  leaflets  nearly  sessile  ;  stem-leaves 

large  and  petioled  :  flowers  bright  lilac,  or  pale  purple,  or  white  ; 

sepals  i  inch  long,  obtuse  ;  limb  of  petal  ^  as  long,  often  white 

in  the  lilac  flowered  form ;  spur  shorter  than  the  limb,  slender 

toward  the  end,  much  incurved  ;  stamens  not  protruding  beyond 

the  petal-limbs  :  follicles  glabrous.      Summer.     Japan.     Revue 

Hort.  109.      1896.     Revue  Belg.  15  :  157. 

Var.  nana-alba  HORT.,  A.  flabellata  v&r.Jlore-albo  HORT. 
Flowers  pure  white  and  the  plant  dwarfish. 

A.  leptoceras  FISCH.  &  MEYER.   Ind.   Hort.   Petrop.  4:  33. 

1837- 

A.  brachyceras  TURCZ.  ex  F.  &  M.   Maund.   Bot.   Gard. 

no.  755.     About  1842. 

Stem  several  flowered,  about  i  ft.  high  :  partial-petioles  of 
root-leaves  over  i  inch,  leaflets  sessile ;  stem-leaves  petioled, 
biternate  :  flowers  violet,  with  the  tips  of  the  sepals  greenish, 
and  tips  of  the  short  petal-limbs  yellow ;  spur  slender,  slightly 
curved,  ^  inch  long,  not  knobbed ;  stamens  protruding  a  little 
beyond  the  limbs  of  the  petals :  follicles  slender,  glabrous, 
nearly  i  inch  long.  Summer.  East  Siberia.  Bot.  Reg.  33  : 
64.  Flor.  des  Serr.  3  :  296.  Little  used  in  America. 

A.  vulgaris  LINN.  Sp.  PL  533.     1753. 

A.  invcrsa  MILL.  Gard.  Diet.  8  ed.  no.  3.      1768. 

A.  silvestris  NECK.  Delic.  Gallo-Belg.  i :   234.      1768. 

A.  cornuta  GILIB.  Fl.  Lituan,  2:  286.     1781. 

A.  versicolor  SALISB.  Prod.  374.     1796. 

A.  corniculata  VILL.  Cat.  Hort.  Strasb.  250.     1807. 

A.  stellata  HORT.  ex  Steud.  Nom.  i  ed.  61.     1821. 

A.  elata  LEDEB.  Ind.  Hort.  Dorp.  Suppl.  41.     1824. 

A.  atrata  KOCH,  in  Flora,  13:   i.     118.     1830. 


338  MINNESOTA    BOTANICAL    STUDIES. 

A.  nigricans  BAUMG.  Enum.  Stirp.  Trans.  2  :  104.    About 

1830. 

A.  Haenkeana  KOCH.  Syn.  Fl.  Germ.  23.      1837. 
A.  concoler  FISCH.  ex  Steud.  Nom.  2  ed.  I  :   115.      1840. 
A.  ecalcarata  HORT.  ex  Steud.  1.  c.  115. 
A.  elegans  POPE  ex  Steud.  1.  c.  115. 
A.  sibirica  DON  ex  Steud.  1.  c.  115. 
A.  subalpina  BOR.  Fl.  Centr.  Fr.  3  ed.  2  :   24.      1840. 
A.  Bernardi  GREN.  &  GODR.  Fl.  Fr.  i  :  45.     1848. 
A.  Transsilvanica   SCHUR.   Vehr.   Siebenb.   Ver.  Nat.  3 : 

94.      1852. 
A.  sylvestris    SCHUR.   Vehr.    Siebenb.  Ver.    Nat.    4:  4. 

1853- 

A.  glaucophylla  STEUD.  in  Flora,  39:  407.      1856. 
A.  aggericola  JORD.  Diagn.  I  :   87.      1860. 
A.  collma]oiLD.  1.  c.  84. 
A.  dumeticola  JORD.  1.  c.  86. 
A.  praecox  JORD.  1.  c.  85. 

A .  paraplesia  SCHUR.  Enum.  PI.  Trans.  28.      1866. 
A.  Corsica  SOLIER.  ex  Nym.  Consp.  18.      1878. 
A  glaucescens  BAKER,  1.  c.  203. 
A.  Karelini  BAKER  1.  c.  76. 
A.  subscaposa  BORHAS  in  Magyar  Tudom.  Akad.  12,  IV, 

18.     1882. 
A.  platysepala  REICHB.    Ic.  Fl.   Germ.   t.  4730.     About 

1886. 

Stem  i  YZ-I  feet  high,  many  flowered ;  finely  pubescent 
throughout :  root-leaves  with  3  partial-petioles  i  ^-2  inches 
long,  secondary  branches  certain,  ultimate  leaf  lobes  shallow 
and  roundish,  texture  firm  ;  lower  stem-leaves  petioled  and  bi- 
ternate  :  flowers  violet  to  dark  purple ;  sepals  ovate,  furnished 
with  a  claw,  acute,  i  inch  long,  one-half  as  wide  ;  petal-limb 
y^  inch  long,  equalling  the  head  of  stamens ;  spur  about  same 
length,  stout,  much  incurved,  knobbed  :  follicles  densely  pubes- 
cent, i  inch  long,  style  half  as  long.  Summer.  Europe, 
Siberia,  and  naturalized  in  America.  Garden  12,  p.  288. 

Var.  nivea  BAUMG.  ex  Baker  Gard.  Chron.  II,  10  :  76.  1878. 
Var.  alba  HORT.  Often  2-3  feet  high  :  a  great  profusion  of 
large,  pure  white  flowers.  Several  weeks  in  early  spring. 

Var.  flore-pleno  HORT.  A.  plena  Hort.  Flowers  much 
doubled,  ranging  from  pure  white  to  deep  blue. 


Davis  :     A    SYXONYMIC    CONSPECTUS    OF    AOJJILEGIAS.  339 

Var.  vervaeneana    HORT.     Var.   folio-aureis  HORT.     Var. 
atroviolacea  HORT.     Leaves  with  yellow  variegated  lines. 
Var.  Olympica  BAKER  1.  c. 

A.  Olympica  BOIES,  in  Ann.  Sc.  Nat.  II,  16 :  360. 
A.  blanda  LEW.  111.  Hort.  4:  t.  14.6.      1857. 
'  A.  Caucasica  LEDEB.  ex  Rupr.  Fl.  Caus.  32.      1869. 

A.   Wittmanniana  STEV.  ex  F.  &  M.  1.  c. 
A  tine  variety  with  several  large  flowers ;   sepals  light  lilac 
or  bright  purple,   i  inch  or  more  in  length,  petal-limb  white. 
Revue  Hort.  1896,  p.  108. 

Var.  hybrida  SIMS.  Bot.  Mag.  1221.     1809. 
Much  like  the  last  variety,  but  with  stout,  lilac-purple  spurs 
as    long    as  the  sepals,   only    slightly    incurved.     Probably  a 
hybrid  of  A.  vulgar  is  and  A.  canadensis. 

A.  Sibirica  LAM.  Encyc.  I  :   150.      1783. 
A.  bicolor  EHRH.  Beitr.  7:   246.      1792. 
A.  speciosa  DC.  Syst.  1 :  336.      1818. 
A.  Garnicriana  SWEET  Brit.    Fl.    Gard.   II,    5:    /.    loj. 

1833- 
Stem  i  *4-2  feet  high,  many  flowered ;  stem  nearly  glabrous 

throughout :  flowers  pale  or  bright  lilac-blue ;  oblong  sepals 
fully  i  inch  long,  spreading  or  reflexed  a  little ;  petal-limb  half 
as  long,  equalling  the  head  of  stamens,  and  often  white ;  spur 
rather  stout,  y2  inch  or  more,  very  much  incurved  or  even 
coiled  :  follicles  glabrous,  i  inch  long,  style  ^  inch.  Allied  to 
A.  vulgaris,  differing  in  the  broad,  obtuse  sepals ;  spur  long 
and  more  slender  toward  the  tip,  and  glabrous  follicles.  Sum- 
mer. East  Siberia.  Sweet's  Brit.  Fl.  Gard.  II,  t.  90  &  t.  loj. 

Var.  spectabilis  BAKER  1.  c.  A.  spectabilis  LEM.  111.  Hort. 
403,  1864.  A  large,  bright  lilac-flowered  variety  with  petal- 
limbs  tipped  with  yellow.  Amurland. 

Var.  jlore-pleno  HORT.  A.  bicolor  var.  jlore-pleno  HORT. 
Flowers  much  doubled  by  the  multiplication  of  both  the  limbs 
and  the  spurs. 

H.  Skinneri   HOOK,  in  Bot.  Mag.  3919.     1842. 
A.  Mexicana  HOOK.,  1.  c. 

Stem  1-2  feet  high,  many  flowered,  glabrous :  root-leaves 
long-petioled,  with  both  primary  and  secondary  divisions  long, 
leaflets  cordate,  3-parted ;  several  stem-leaves  petioled  and  bi- 
ternate  :  sepals,  green,  keeled,  lanceolate,  acute,  never  much 


340  MINNESOTA    BOTANICAL    STUDIES. 

spreading,  ^-i  inch  long ;  petal-limb  greenish-orange,  half  as 
long  as  sepal ;  spur  bright  red,  tapering  rapidly,  over  i  inch 
long ;  stamens  protruding  far  beyond  the  limb  :  fruit,  at  least 
when  young,  bearing  broad,  membranous,  curled  wings  ;  styles 
3 ;  after  flowering  the  peduncles  become  erect.  July-Sept. 
Mts.  of  New  Mex.  andMex.  Bot.  Mag.  3919.  Paxt.  Mag.  Bot. 
10  :  199.  Flor.  des  Serr.  i  :  17.  A  handsome  plant,  requiring 
a  light  soil  in  a  sunny  border. 

Var.  Jlore-pleno  HORT.  Flowers  double.  Very  fine.  Gar- 
tenflora  34:  57. 

A.  formosa  FISCH.  in  DC.  Prod,  i :  50.     1824. 
A.  artica  LOUD.  Hort.  Brit.  610.      1830. 
A.   Canadensis  var.  formosa  WATS.     Bot.  King  Exp.  10. 

1871. 

Habit  as  A.  Canadensis,  root-leaves  and  stem-leaves  like  that 
species,  but  the  flowers  are  brick  red  and  yellow  or  wholly  yel- 
low, and  the  sepals  are  larger,  twice  as  long  as  the  petal-limb, 
more  spreading ;  spurs  somewhat  more  slender  and  often 
shorter.  May- Aug.  Sitka  to  Calif,  and  eastward  to  the  Rock- 
ies. Introduced  1881.  Bot.  Mag.  6552.  Flor  des  Serr.  8: 

795- 

Var.  desertorum  JONES  Cont.  Western.  Bot.  No.  8,  2.   1898. 

Stems  about  one  foot  high,  flexuose :  leaves  and  leaflets 
rather  small :  flowers  often  only  about  ^  inch  across,  nodding  : 
styles  slender.  In  crevices  of  rocks  about  springs.  Flagstaff, 
Ariz.  (f). 

Var.  truncata  BAKER  1.  c. ;  JONES  Zoe,  4:  259.     1893. 
A.  truncata  FISCH.  &  MEY.      1843. 
A.  California*  LINDL.  in  Gard.  Chron.  836.      1854. 
A.  eximia  VANHOUTTE  ex  Planch.  Fl.  des  Serr.  12  :   1188. 

1857- 

Flowers  with  short  thick  spurs  and  very  small  sepals  and  petal- 
limbs.  Introduced  1881. 

Var.  hybrida  HORT.  A.  Californica  var.  hybrida  HORT. 
Flowers  large  with  scarlet  sepals  and  yellow  petals ;  spurs 
spreading,  long  and  slender.  A  supposed  hybrid  with  A. 
chrysantha.  Fl.  Mag.  1877  :  278.  Vicks'  i  :  33,_/".  2. 

Var.  nana-alba  HORT.  Flowers  pale,  often  nearly  white  ; 
plant  not  exceeding  i  foot. 

Var.  rubra-pleno  HORT.  Flowers  as  in  var.  hybrida,  but  with 
several  whorls  of  petal-limbs. 


Davis  :   A  SYNONYMIC  CONSPECTUS  OF  AQUILEGIAS.         341 

A.  pubescens  COVILLE  Contr.  Nat.  Herb.  4:  56,  _^>/.  /.    1893. 

Allied  to  A.  chrysantha.  Caudex  scaly;  flowers  a  very 
clear  yellow  ;  spurs  shorter,  only  i  to  i  ^  inches  ;  petal-limbs  a 
third  as  long  as  sepals.  July.  High  altitudes  Tulare  Co.,  Calif. 

A.  chrysantha  GRAY  in  Gard.  Chron.  1335  and  1501,  f.  304. 

1873- 

A.  leptoceras  var.  chrysantha  HOOK.  f.  Bot.   Mag.  6073. 


A.  leptoceras  var.jZava  GRAY  PL  Wright  2  :  9.     1852. 

Height  3-4  feet;  root-leaves  with  twice  3-branched  petioles, 
leaflets  biternate  ;  stem-leaves  several,  petioled  ;  flowers  many 
on  the  plant  ;  2-3  inches  across  ;  sepals  pale  yellow,  tinted 
claret,  spreading  horizontally  ;  petal-limbs  deep  yellow,  shorter 
than  the  sepals  and  nearly  as  long  as  the  head  of  stamens  ; 
spur  rather  straight,  very  slender,  divergent,  about  2  inches 
long,  descending  when  flower  is  mature  ;  follicles  glabrous,  I 
inch  long  ;  style  half  as  long.  May-  Aug.  New  Mex.  and 
Ariz.  Introduced  1891.  Revue  Hort.  1896:  108.  Flor.  des 
Serr.  20;  2108.  Floral  Mag.  1873  :  88.  Diet.  d.  Bot.  i  :  243. 
Vicks'  i  :  33,  /.  3. 

Var.  aurea  n.  var. 

A.  aurea  JUNKA.  in  Oestr.  Bot.  Zeitschr.  22:   174.      1872. 
A.  Canad^ensis  var.  aurea  ROEZL.,  Gartenflora  258,  t.  734. 
1872. 

Flowers  yellow  and  tinged  with  red,  spurs  incurved  and 
shorter  than  in  the  type.  Gartenflora  21  :  734. 

Var.  alba-plena  HORT.  Var.  grandiflora-alba  HORT.  Flowers 
very  pale  yellow  or  nearly  white,  with  two  or  more  whorls  of 
petal-limbs.  Introduced.  1889.  Vicks'  12  :  311. 

Var.  nana  HORT.  A.  leptoceras  var  lutea  HORT.  Like  the 
type,  but  plant  always  small,  not  exceeding  i^£  feet. 

Var.  Jaeschkani  HORT.  About  the  same  height  as  the  last  : 
flowers  large,  yellow  with  red  spurs.  Thought  to  be  a  hybrid  of 
A.  chrysantha  and  A.  Skinneri,  hence  sometimes  called  A. 
Skinneri  var.  hybrida  HORT. 

A.  longissima  GRAY  ex  Wats.  Proc.  Am.  Acad.  17:  317. 
1881-2. 

Tall,  somewhat  pubescent  with  silky  hairs,  or  smoothish  : 
root-leaves  biternate  even  in  the  petioles,  leaflets  deeply  lobed 
and  cut,  green  above,  glaucous  beneath  ;  stem-leaves  similar, 


342  MINNESOTA    BOTANICAL    STUDIES. 

petioled  :  flowers  pale  yellow,  sepals  lanceolate,  broadly  spread- 
ing, an  inch  or  more,  the  spatulate  petals  a  little  shorter,  about 
equalling  the  head  of  stamens  ;  spur  4  inches  long  or  more,  al- 
ways hanging,  orifice  narrow.  Distinguished  from  A.  chrysan- 
tha  by  longer  spur  with  contracted  orifice,  by  the  narrow  petals, 
and  by  the  late  season  of  flowering.  Late  July  to  October  ist. 
Ravines  southwest  Texas,  into  Mexico.  The  seed  must  be  ob- 
tained from  the  wild  plants,  as  those  cultivated  usually  fail  to 
produce  seed;  hence  not  much  used.  Garden  &  Forest  i  :  31. 

A.  caerulea  JAMES,  Long  Exped.  Rocky  Mts.  2:  15.     1826. 
A.  Zeptocera  NUTT.  in  Journ.  Acad.  Phila.  7:  9.      1834. 
A.  macrantha  HOOK.  &  ARN.  Bot.  Beech.  Voy.  317,  t.  72. 
1841. 

Stem  i^  feet  high,  finely  pubescent  above,  bearing  several 
flowers  :  lower  stem-leaves  large  and  biternate ;  basal  leaves 
with  long  3-branched  petioles ;  leaflets  3-lobed  on  secondary 
stalks  :  flowers  2  inches  across,  whitish,  but  variously  tinted 
with  light  blue  and  yellow ;  sepals  often  blue,  oblong,  obtuse, 
twice  as  long  as  the  petal-limbs  :  spurs  long,  slender,  knobbed 
at  the  end,  rather  straight  but  curving  outward  :  head  of  stamens 
equalling  the  petals  :  follicles  pubescent,  i  inch  long ;  style  % 
inch.  April-July.  Lower  mountain  regions,  Montana  to  New 
Mexico.  Introduced  1891.  Bot.  Mag.  4407.  Garden  16 : 
198.  Meehan's  Mo.  6 :  61.  Vicks'  i:  33,  f.  4.'.  Bot.  Mag. 
5477  (as  var.  ochroleuca).  Flor.  des  Serr.  5  :  531. 

Var.  albiflora  GRAY  Syn.  Fl.  i :    i  :  44.     1895. 

Flowers  of  the  same  size  but  sepals  as  wrell  as  petals  almost 
white.  Introduced  as  var.  alba  HORT.  1883. 

Var.  alpina  A.  NELSON  Wyom.  Exp.  Sta.  Bull.  28 :  78. 
1895-6. 

Plant  smaller  than  the  type  ;  upper  leaflets  entire ;  flowers 
smaller,  yellow,  spurs  rather  short. 

Var.  calcarea  JONES  Proc.  Calif.  Acad.  Sci.  II.  5 :  619. 
1895. 

Plant  glandular-pubescent :  leaves  and  leaflets  reduced,  firm  ; 
flowers  much  smaller  than  the  type  ;  sepals  blue-purple  ;  petals 
reddish.  Cannonville,  Utah. 

Var.  hybrida  HORT.  Sepals  some  shade  of  blue  or  pink  or 
mixed,  and  petals  nearly  white  or  yellow.  The  true  form  of 
this  is  probably  A.  ccerulea  X  A.  chrysantha.  Revue  Hort. 
1896:  108.  Am.  Gard.  15:  315. 


Davis:   A  SYNONYMIC  CONSPECTUS  OF  AOJJILEGIAS.         343 

Var.  flore-pleno  HORT.     Flowers  longer  and  very  showy ; 
more  or  less  doubled  toward  the  center. 

A.  alpina  LINN.  Sp.  PI.  533.     1753. 

A.  montana  STERNB.  in  Regensb.  Denkschr.  2:  60.    1818. 

A.  Reutcriana  REICHB.  f.  Nym.  Consp.  18.      1878. 

A.  Sternbergii  REICHB.  Fl.  Germ.  Excurs.    749.     About 
1880. 

A.  alpina  var.  superba  HORT. 

Stem  nearly  i  ft.  high,  finely  pubescent  in  upper  parts  ;  2-5 
flowered:  leaves  biternate,  petioled ;  partial-petioles  of  basal 
leaves  1-2  inches  long  with  3  nearly  sessile  leaflets,  deeply 
lobed :  expanded  flower  1*^-2  inches  across,  blue,  rarely  pale 
or  white  :  sepals  i%  inches  long,  half  as  broad,  acute;  petal- 
limb  half  as  long  as  sepal,  often  white ;  spur  slender,  incurved, 
same  length  as  the  limb  :  head  of  stamens  not  protruding  :  fol- 
licles pubescent,  i  inch  long,  style  much  shorter.  May-June. 
Switzerland.  Bot.  Cab.  7:  657.  Garden  9  :  17. 

A.  glandulosa  FISCH.  Hort.  Gorenk.  2ed.  48.     1812. 

Stem  i  to  i  y?  feet,  high ;  glandular-pubescent  in  the  upper 
half;  1-3  flowered:  partial-petioles  of  root  leaves  1-2  inches 
long,  each  with  3  distinct  divisions ;  leaflet  segments  narrow 
and  deep  :  stem-leaves  few,  bract-like  :  flowers  large,  nodding  : 
sepals  bright  lilac-blue,  ovate,  acute,  almost  i^  inches  long 
and  half  as  broad ;  petal-limb  same  color,  but  tipped  and  bor- 
dered with-creamy  white,  less  than  half  the  length  of  the  sepals, 
very  broad ;  spur  very  short,  ^  inch,  stout,  much  incurved ; 
stamens  not  protruding ;  follicles  i  inch  long,  6-10  in  number, 
densely  hairy,  with  short  falcate  style.  Allied  to  A.  alpina,  but 
with  shorter  spurs,  larger  flowers,  plant  taller,  greater  number 
of  follicles.  May-June.  Altai  Mts.  of  Siberia.  Botanist  5:  219. 
Floral  World  1871  :  354.  Garden  15  :  174.  Gartenflora  289,  f.  L 
Var.  jucunda  FISCH.  &  LALL.  Ind.  Sem.  Petrop.  2.  1840. 
Flowers  rather  smaller  than  in  the  type ;  petal-limb  white,  more 
truncate  at  the  tip ;  stamens  as  long  as  the  limb.  A  variety 
with  some  tendency  to  double.  Bot.  Reg.  33  :  19.  Flor.  des 
Serr.  5:  535. 

A.  Stuarti  HORT.  A  recorded  hybrid  of  A.  glandulosa  and 
A.  vulgaris  var.  Olympica.  Flowers  very  much  resemble  the 
latter  parent  in  form  of  sepals  and  petals,  and  the  former  in 
shape  of  spurs  and  in  coloration.  May-June.  Introduced 
1891.  Garden  34  :  670. 


XXI.     A   SYNONYMIC   CONSPECTUS   OF  THE   NA- 
TIVE  AND   GARDEN   ACONITUMS    OF 
NORTH   AMERICA. 


K.  C.  DAVIS. 


This  genus  of  hardy,  ornamental,  perennial  herbs  is  much 
used  in  borders,  and  such  places,  and  is  commonly  called 
Monkshood.  Many  species  are  planted  in  European  gardens ; 
nine  only  have  been  thus  used  in  America.  The  number  of 
species  varies  from  18  to  80,  according  to  treatment  by  differ- 
ent authors.  They  are  native  in  mountainous  regions  of 
Europe,  temperate  Asia,  and  7  are  found  in  North  America. 

Roots  tuberous,  turnip-shaped,  or  thick-fibrous  :  stem  tall  or 
long,  erect,  ascending,  or  trailing :  leaves  palmately  divided 
or  cleft  and  cut-lobed  :  flowers  large,  irregular,  showy ;  sepals 
5,  the  large  upper  sepal  in  shape  of  a  hood  or  helmet;  petals 
2-5,  small;  stamens  numerous;  carpels  3-5,  sessile,  many- 
ovuled,  forming  follicles  when  ripened. 

The  following  species  do  well  in  any  garden  soil,  but  rich  soil 
is  preferred.  They  thrive  in  open  sun,  but  flowers  last  longer 
in  shaded  places.  Aconites  should  never  be  planted  in,  or  too 
near  the  kitchen  garden,  or  the  children's  garden,  as  the  roots 
and  some  of  the  flowers  contain  a  deadly  poison.  Propagated 
easily  by  root  division. 

Besides  the  Prodromus  treatment  by  A.  DeCandolle,  in  1824, 
the  only  other  monographs  of  the  whole  genus  are  by  H.  G. 
L.  Reichenbach,  "  Uebersicht  der  Gattung  Aconitum,"  Leip- 
sic,  1819;  "  Monograph  Generis  Aconiti,"  Leipsic,  1820,  folio, 
2  vols.,  and  "  Illustratio  Spec.  Aconiti,"  Leipsic,  1823-7,  folio. 
Reichenbach  considered  the  number  of  species  to  be  about  80, 
but  many  of  his  names  should  be  treated  as  synonyms,  as  they 
were  given  to  forms  varying  only  slightly  from  the  type. 


346  MINNESOTA  BOTANICAL  STUDIES. 

KEY  TO  SPECIES  OF  ACONITUM. 

A.   Roots  tuberous  or  napiform. 

B.  Leaves  deeply  cut,  but  not  to  the  base. 
C.   Sepals  mostly  some  shade  of  blue. 

D.   Axils  of  leaves  without  bulblets. 

E.   Beak  of  helmet  prominent  or  reflexed. 

F.   End  of  beak  reflexed Cammarum. 

FF.  End  of  beak  not  reflexed,  but  prominent. 

G.   Stem  stout paniculatitm. 

GG.   Stem  lax K amis  chat  icum. 

EE.   Beak  of    helmet   either    abruptly  pointed    or 
turned  inward — not  reflexed. 

F.   Stem  stout,  erect .Japonicum. 

FF.   Stem  slender,  sometimes  showing  a  climb- 
ing tendency nncinatum. 

DD.   Axils  of  leaves  with  bulblets bulbiferum. 

CC.   Sepals  mostly  white  or  yellow  bordered  with  blue. 

D.   Stem  robust heterophyllum. 

DD.   Stem  slender Lycoctonum. 

BB.   Leaves  divided  to  the  base. 

C.  Helmet  higher  than  wide variegatunt. 

CC.  Helmet,  or  hood,  broad  and  low. 

D.   Follicles  rarely  varying  from  four Napellus. 

DD.   Follicles  three  to  five delphinifolium. 

AA.   Roots  fascicled  and  elongated,  or  fibrous. 
B.   Stems  erect. 

C.   Flowers  yellowish Anthora. 

CC.   Flowers  blue  to  whitish autumnale. 

BB.   Stems  trailing reclinatum, 

A.  cammarum  LINN.  Sp.  PI.  2  ed.  751.     1762. 

A.  neomontanum  WILLD.  Sp.  PL  2:   1236.      1799. 

A.  bicolor  SCHULT.  Obs.  Bot.  101.      1809. 

A.  erioslemum  DC.  Syst.  I:   377.      1818. 

A.  intermedium  DC.  1.  c.  374. 

A.  spectosum  OTTO,  ex  DC.  Prod.  1 :  60.      1824. 

A.  cernuum  BAUMG.  ex.    Schur.   Enum.   PL   Transs.   32. 

1866. 

REICHENBACH  has  given  the  names:  atistriacum  TRATT., 
Breiterianum,  decorum,  exaltatum,  hamatum,  hortense 
HOPPE.,  Ottonianum,  -palmatifidum,  Sprengelii  RUA, 
Storkianum,  versicolor. 

Stem  3-4  feet  high  ;  leaves  with  short  bluntish  lobes  ;  flowers 
purple  or  blue  ;  panicles  or  loose  spikes  few  flowered  ;  helmet 


Davis:   A  SYNONYMIC  CONSPECTUS  OF  ACONITUMS.         347 

hemispherical,  closed ;  beak  reflexed.  July-Sept.  Hungary. 
Introduced  1889.  A.  Storkianum  RCHB.  Uebers  49,  is  a  form 
of  this  with  dwarf  habit  and  fewer  flowers;  roots  somewhat 
fibrous.  Bot.  Cab.  1991. 

A.  paniculatum  LAM.  Fl.  Fr.  3:  646.     1778. 

A.  cernuum  WULF.  ex  Koelle,  Spicil.  17.     1786. 
A.  humile  SALISB.  Prod.  375.     1796. 
A.   Wilematianum  DELARB.  Fl.  Anv.  2  ed.  499.      1800. 
A.  hebcgynum  DC.  Syst.  i  :  376.     1818. 
A.  camarum  SCHLEICH.  Cat.  5.     1821. 
A.  neomontanumBAUMG.  Enum.  Transs.  2  :  100.   1816-46. 
Forms  were  named  by  REICHENBACH  :  acuminatum,  plexi- 
canle  HOPPE  &  HORN,  galectonum,  gibbiferum,  molle, 
parviflorum,  taxicum,  reflexum. 

Stem  erect,  slender,  2-3  feet  high ;  leaves  glabrous,  rather 
thin,  deeply  3-7  cleft,  the  divisions  obovate,  cuneate,  incised 
and  dentate ;  inflorescence  pubescent,  panicled ;  flowers  blue, 
few,  half  as  broad  as  high ;  helmet  with  a  gibbous  arch ;  beak 
very  prominent  and  descending,  %  inch  long ;  open ;  follicles 
usually  4,  erect,  %  inch  long.  Summer.  Central  Europe. 
Well  figured  in  Bot.  Cab.  810. 

A.  Noveboracense  Gray  ex  Coville  in  Bull.  Torr.  Club,  13 : 
190.  1886,  of  South-eastern  New  York,  differs  very  little,  if 
any,  from  this. 

A.  Kamtschaticum  PALL,  ex  Rchb.  Uebers.  39.     1819. 
A.  Nafettus  THUNB.  Fl.  Jap.  251.     1784,  not  Linn. 
A.  maximum'?  KLL.  ex.  DC.   Syst.  i:  380.     1818  (name 
only). 

A.  Ftschert"R.cm.  Monog.  t.  22.     1820. 

A.  Labanskyi  RCHB.  1.  c.  t.  40. 

A.  abbreviation  LANGSD.  ex  DC.  Prod,  i :  61.     1824. 

A.  nasutum  HOOK.  Fl.  Bor.  Am.  i :   26.     1833. 

A.  sinense  SIEB.  &  Zucc.  1835,  not  Chinensis  Siebold. 

A.  Columbianum    NUTT.   in  Torr.    &  Grav,   Fl     i  •    i* 

1838. 

A.  autumnale'LiND.  in  Journ.  Hort.  Soc.  2:   77.      1847. 
A.  arcuatum  MAXIM.  Fl.  Amur.  27.     1859. 
A.  Californicum  Hort. 

Roots  napiform ;  stem  lax,  2-4  ft.  high,  pubescent  in  upper 
parts  :  leaves  parted  or  deeply  cleft,  divisions  broadly  ovate  or 


348  MINNESOTA    BOTANICAL    STUDIES. 

cuneate  and  incised  and  dentate :  flowers  pale  blue ;  helmet 
higher  than  broad,  being  %-^  in.  long  ;  beak  often  elongated ; 
lower  petals  small,  oblong :  follicles  erect,  oblong,  reticulate. 
Autumn.  Of  wide  geographic  range,  being  found  in  many 
parts  of  Asia,  Europe  and  in  the  U.  S.  west  of  the 
Introduced  1889.  Bot.  Mag.  7130. 

A.  Japonicum  DECNE.  in  Rev.  Hort.  473.     1851. 

Stem  erect,  3-4  ft.  high,  smooth ;  leaves  dark  green,  shining, 
petioled,  lobes  2-3  times  cut,  parts  blunt  and  deeply  toothed ; 
flowers  large,  deep  blue  or  violet  tinged  with  red;  loose  pan- 
icles with  ascending  branches ;  helmet  conical,  beak  abruptly 
pointed:  follicles  5.  July-Sept.  Japan.  Introduced  1889. 

Rev.  Hort.  1.  c. 

Var.   coeruleum    Hort.      Flowers    very    abundant;  panicle 

shortened. 

A.  uncinatum  LINN.  Sp.  PI.  2  ed.  750.     1762. 
A.  Japonicum  THUNB.  Fl.  Jap.  232.      1784. 
A.  volubile  MUHL.  Cat.  52.     1813. 
A.  scandens  MUHL.  ex  Rchb.  Uebers.  38.     1819. 
A.  -variegatum  HOOK.  f.  &  THOMS.  Fl.  Ind:  56.     1858. 
Stem  slender,  3-5  ft.  high;  inclined  to  climb  ;  glabrous  below 
the  inflorescence  :  leaves  thick,  deeply  cut  into  3-5  cut-toot! 
lobes  :  flowers  loosely  panicled  but  crowded  at  the  apex  ;  blue, 
pubescent,  i  inch  broad ;  helmet  erect,  nearly  as  broad  as  long, 
obtusely  conic,  beak  or  point  of  helmet  turned  inward :  follicles 
3,  y,  in.  long.     June-Sept.     Low  grounds  of  Penn.,  South  a 
West,  Japan.    Much  planted  now.    Meehan's  Mo.  4  :  81.     Bot. 
Mag.  1119. 

A.  bulbiferum  HOWELL,  Fl.  N.  W.  Am.  1 :  25.     1897. 
Stems  slender,  weak  and  viney,  2-4  ft.  long  ;  smooth  belo\i 
tomentose  above  :  leaves  rather  small,  on  short  petioles,  or  t 
upper  sessile,  bearing  bulblets  in  their  axils,  all  laciniately  cu 
into  acute  lobes  :  sepals  pale  blue  ;  hood  6-8  lines  long, 
not  seen.     In  marshes  on  the  eastern  slope  of  the  Cascade  Mti 
near    Mt.    Hood.     Flowering    in    Sept.     Described   from   the 
original. (f) 

A.  heterophyllum  WALL.  Cat.  4722.    1828.    Bot.  Mag.  6092. 

1874.  " 

A.  cordatum  ROYLE.     Illustr.     56.     1839.  not  Rafin. 
Stem  robust,  3  ft.    high :  lower  leaves  petioled,  upper  one 


Davis :   A  SYNONYMIC  CONSPECTUS  OF  ACONITUMS.         349 

sessile  ;  all  dark,  large,  cordate,  coarsely  dentate  :  sepals  yellow 
with  violet  or  blue  margins ;  hood  arched,  beak  rather  blunt : 
follicles  5,  erect  pubescent.  Widely  distributed.  Himalaya  re- 
gion. Not  yet  introduced  to  the  American  trade,  but  has  re- 
cently been  used  in  European  gardens  because  of  its  striking 
flowers  and  leaves.  It  is  used  in  India  as  a  tonic  medicine. 

A.  Lycoctonum  LINN.  Sp.  PI.  532.     1753. 
A.  Pyrenaicum  LINN.  1.  c.  532. 
A.  altissimum  Mill.  Gard.  Diet.  8  ed.  no.  2.      1768. 
A.  Napellus  S.  G.  GMEL.  It  1 :  8.      1768  (?). 
A.  septentrionalc  KOELLE.  Spicil.  22.      1786. 
A.  taxicarium  SALISB.  Prod.  375.      1796. 
A.  intermedeum  Host.  Fl.  Aust.  2:  69.      1797. 
A.  Jacquinianum  HOST.  1.  c.  68. 
A.  pauciflorum  HOST.  1.  c.  70. 
A.  ochroleucum  WILLD.  Sp.  PL  2:  1233.     1799. 
A.  barbatum  PATR.  ex  Pers.  Syn.  2  :  83.     1807. 
A.  galeriflorum  STOKES,  Bot.  Mag.  Med.  3:  216.     1812. 
A.  Ncapolitanum  TENORE,  Fl.  Nap.  1 :  327.     1815. 
A.  hispidiim  DC.  Syst.  I  :  367.      1818. 
A.  squarrosum  LINN,  ex  DC.  1.  c.  368. 
A.  ochranthum  C.  A.  MEY.   in  Ledeb.  Fl.  Alt.   2:   285. 

1830. 
A.    delphinifolium    HORT.   ex   Steud.  Nom.  2"    ed.   I  :   18. 

1840. 

A.  ochroleucum  HORT.  ex  Steud.  1.  c.  19. 
A.  rubicundum  FISCH.  ex  Steud.  1.  c.  20. 
A.  triste  FISCH.  ex  Steud.  1.  c.  20. 
A.  excelsum  TURCZ.  Cat.  Baikal.  70.      1842. 
A.  Hosteanum  SCHUR.  in  Verh.  Seibenb.  Ver.  Naturw.  2: 

177.      1851. 
A.  Transilvanicum  LERCH.  ex  Schur.  in  same,  10  :  165. 

1859. 
Lycoctonum  sylvaticum  FOURR.  in  Ann.  Soc.  Linn.  Lyon, 

n.  s.  16:   326.      1868. 
A.  umbracticolum  SCHUR.  in  Verh.   Naturf.  Ver.  Bruenn. 

15:    2.        1877. 

Names  of  this  accredited  to  REICHENBACH  are :  cego- 
phonum,  alienum,  arctophonum,  australe,  loreale  SER., 
cynoctonum,  dissectum  TAUSCH.,  Gmelini,  lagoctonum, 
Lamarckii,  luparia,  lupicida,  meloctonum,  moldavicum 


350  MINNESOTA    BOTANICAL    STUDIES. 


monanense    SCHMIDT.,    myoctonum,    -pallidum, 
Phthora,  ranunculi  folium,  rectum  BERNH., 
strictissimum,    strictum   WILLD.,    thelyphonum^   therio- 
•phonum,  tragoctonum,  vulparia,  zooctonum. 
Stem  slender,  simple,  3-6  feet  high;  leaves  deeply  cut  into 
5—9  lobes  ;  long  petioles  and  under  ribs  pubescent  ;  flowers  yel- 
low or  whitish  in  racemes  ;   helmet  a  pinched  elongated  cone  ; 
middle  sepals  usually  bearded  :  follicles  usually  3.    June-Sept. 
Europe;   Siberia.     Bot.  Mag.  2570.      Gard.  Mag.  34:    124. 

A.  variegatum  LINN.  Sp.  PL  532.     1753. 

A.  alpinum  MILL.  Gard.  Diet.  8  ed.     No.  7.      1768. 

A.  cammarum  JACQ^  Fl.  Aust.  5.  /.  224.     1775. 

A.  luridum  SALISB.  Prod.  375.      1796. 

A.  volubile  MOENCH.  Meth.  Suppl.  ITO.     1802. 

A.  ro  stratum  BERNH.  Ind.  Sem.  Hort.  Erf.      1815. 

A.  glabrum  DC.  Syst.  I  :   379.      1818. 

A.  leucanthemum  WENDER.  in  Linnaea  5:   53.      1830. 

A.  nasutum  FISCH.  ex.  G.  Don.  Gen.  Syst.  I  :  61.     1831. 

A.  intermedium  GAUD,  ex  Steud.  Nom.  2  ed.  I  :  18.    1640. 

A.  Japonicum  Hort.  ex  STEUD.  1.  c.  18. 

A.  laciniosum  SCHLEICH.  ex  Steud.  1.  c.  18. 

A.  laevigatum  SCHLEICH.  ex  Steud.  1.  c.  18. 

A.  uncinatum  Hort.  ex  STEUD.  1.  c.  20. 

A.  altigaleatum  HAYNE,  Arzn.   Gew.  12,  /.  16.      1845. 

A.  Burnhardianum  WALLR.   Sched.    Crit.    I  :    250,  /.  2. 

1848. 

REICHENBACH'S  names  for  forms  of  this  are  :  bulbiferum, 
flexuosum  PRESL.,  gracile,  hamatum,  illinitum,  italicum 
TRATT.,  lasiocarpum,  macranthum,  mixtum,  rhynchan- 


Erect,  1-6  ft.  high  :  leaves  variously  divided  into  usually 
broad  lobes  and  cut  divisions  ;  lower  petioles  long,  others  short 
or  none  :  flowers  in  a  loose  panicle  or  raceme  ;  blue  varying  to 
whitish,  smoothish  ;  helmet  higher  than  wide,  top  curved  for- 
ward ;  beak  pointed  horizontal  or  ascending.  July.  Europe. 
Var.  album  n.  var. 

A.  album  AIT.  Hort.  Kew.  2  :   246.     1789. 
A  pure  white  form  of  above  type,  with  roots  rather  fibrous. 

A.  Napellus  LINN.  Sp.  PI.  532.     1753. 

A.  -pyramidale  MILL.  Gard.  Diet.  8  ed.  no.  6.      1768. 


Davis:   A  SYNONYMIC  CONSPECTUS  OF  ACONITUMS.         351 

A.  lauricum  WULF.  in  Jacq.  Coll.  2:   112.      1788. 

A.  neubergense  DC.  Syst.  I  :  373.      1818. 

A.  strictum  BERNII.  ex  DC.  1.  c.  373. 

A.  laxiflornm  SCHL.  Cat.  5.      1821. 

Napellus  vulgar  is  FOURR.   in  Ann.  Soc.  Linn.  Lyon,  n. 

s.  16:   326.      1868. 

REICHENBACH  has  given  twenty-four  names  to  forms  of 
this :  acutum,  ambiguum,  amcenum,  Bcrnhardianum, 
Braunii)  callibotryon,  clusianum,  commutatum,firmum, 
formosum,  Funkianum,  /lians,  Hoppeanum,  Koehleri, 
IcBtum,  laxiflorum  SCHL.,  laxum,  Mielichhoferi,  napel- 
loides  Sw.,  oligocarpum,  rigidum,  tennifolium,  venus- 
tum,  virgatum. 

The  best  known,  as  well    as    the   most  poisonous   species. 
Stem  erect,  3-4  ft.  high  :  leaves  divided  to  the  base  and  cleft 
—3  times  into  linear  lobes  :  flowers  blue  in  a  raceme  :  peduncles 
rect,  pubescent :   helmet  broad  and  low,  gaping,  smoothish  : 
:ollicles    4,    rarely  3.     June-July.      Europe,    and   naturalized 
icre.       Gartenflora,   35  :     227    (named    A.    dissectum    Don.), 
iegne  Vegetal,  Med.  PI.  2.     There  are  very  many  varieties, 
iffering  in  shade  of  flowers,  often  mottled  or  lined  with  white. 
Var.  album  is  nearly  white.     Var.  bicolor  and  Var.  versicolor 
are  much  used  in  gardens  for  the  large  blue-and-white  flowers. 
Jot.  Cab.  8  :   794  (vcrsicolor}. 

A.  delphinifolium  DC.  Syst.  1 :  380.     1818. 

A.  Napellus  var.  delphinifolium  SERINGE,  Mus.  Helv.  i : 

159.     1823. 
REICHENBACH   used  for  forms  of  this  :    Chamissonianum, 

paradoxum,  semigaleatum. 

Stem  erect,  1-3  feet  high,  pubescent :  leaves  deeply  parted, 
ind  cleft  into  narrow  lobes :  flowers  blue,  large ;  hood  low, 
>eak  short ;  lateral  sepals  as  long  and  twice  as  broad  as  the 
ower  :  follicles  oblong.  British  Columbia  and  Alaska  through 
slands  of  Behring  Strait  to  Asia. 

Var.  ramosum  n.  var. 

A.    ramosum  A.  NELS.  Bull.  Torr.  Club  26:  8.      1899. 
Much  like   the  type  :  leaves  larger  with  fewer  and  longer 
egments  :  follicles  less  pubescent.     Professor  Nelson   consid- 
ers the   pubescence   different.      Only  once  found :    Limestone 
iange,  Black  Hills,  Weston  Co.,  Wyom.     Co-type  at  Colum- 


352  MINNESOTA    BOTANICAL    STUDIES. 

A.  Anthora  LINN.  Sp.  PL  532.     1753. 

A.  Pyrenaictim  PALL.   Reise.  2  :   316.      1776. 

A.  ochroleucum  SALISB.  Prod.  375.      1796. 

A.  Anthorideum  DC.  Syst.  I  :  366.      1818. 

A.  Anthorum  ST.  LAG.  in  Ann.  Soc.  Bot.  Lyon,  7:   119. 
1880. 

REICHENBACH   used    these   names :     Candollei^   eulophum, 
Jacquini,  nemerosum,  BIEB.,  Pallasii,  tuberosum  PATR. 

Stem  erect,  1-2  feet  high  :  leaves  parted,  usually  at  the  base, 
parts  deeply  cut  and  lobed  ;  more  or  less  hispid  beneath,  smooth- 
ish  above ;  petioles  long :  flowers  in  lateral  and  terminal 
racemes,  pale  yellow,  often  large  ;  inflorescence  generally  pu- 
bescent ;  spur  refracted  or  hooked  ;  helmet  arched  but  cylindri- 
cal at  base  :  follicles  5.  June-July.  Southern  Europe.  There 
are  several  garden  varieties  differing  in  pubescence,  size  of 
flower,  shape  of  galea,  and  width  of  leaf  segments.  Bot.  Mag. 
2654. 

A.  autumnale  REICHB.  Monog.  t.  //,_/".  2.     1820. 

Stem  erect,  3-5  feet  high :  leaves  pedately  five  lobed : 
flowers  in  simple  spike  becoming  a  panicle ;  blue,  lilac,  or 
whitish ;  helmet  closed.  Sept. -Nov.  North  China.  Intro- 
duced about  1870. 

A.  reclinatum  GRAY  Am.  Journ.  Sci.  42  :  34.      1842. 

Stem  always  trailing,  2-5  feet  long,  nearly  glabrous ;  leaves 
thin,  deeply  3-7  cleft,  toothed  and  cut,  lower  ones  petioled, 
large,  upper  ones  sessile ;  flowers  white  or  dull  cream-color, 
pubescent,  in  loose  raceme  or  simple  panicle  :  helmet  twice  as 
high  as  wide,  conic;  beak  very  short;  follicles  3,  nearly  ^ 
inch  long.  Summer.  Wooded  mountain  regions  of  Va.  to 
Ga.  (f). 

NOTE  :  The  mark  (t)  indicates  that  the  native  species  or  variety  has  not  jet 
been  introduced  to  the  American  trade.  Citations  at  the  end  of  a  description 
are  mostly  to  colored  plates. 


XXII.     A   CONTRIBUTION    TO    THE    KNOWLEDGE 

OF   THE   FLORA  OF   SOUTHEASTERN 

MINNESOTA. 


W.  A.  WHEELER. 


The  work  of  the  Minnesota  Botanical  Survey  in  southeastern 
Minnesota  during  the  summer  of  1899  was  carried  on  with  two 
main  purposes  in  view  :  first,  to  collect  and  preserve  plants  in 
formalin  for  museum  and  class  use,  and  second,  to  collect 
herbarium  specimens  of  the  higher  seed  plants.  The  work  of 
collection  was  begun  June  ist,  and  closed  August  3ist.  The 
catalogue  of  species  is,  therefore,  very  incomplete  in  its  enu- 
meration of  the  early  spring  and  autumn  plants. 

District  of  collection. — The  territory  in  which  the  collections 
were  made  is  in  the  extreme  southeastern  part  of  Minnesota, 
comprising  the  valleys  of  Winnebago  and  Crooked  creeks,  and 
the  adjoining  region  near  the  Mississippi  river.  Nearly  all  of 
this  territory  is  included  in  an  area  about  twelve  miles  square, 
formed  by  the  townships  of  Mayville  and  Crooked  Creek,  on 
the  north,  and  Winnebago  and  Jefferson  on  the  south. 

Physiography. — The  topography  of  this  part  of  Houston 
county  is  not  essentially  different  from  that  of  most  of  the  re- 
gion south  from  Red  Wing  along  the  Mississippi  river  to  the 
southern  boundary  of  Minnesota  and  into  Iowa.  There  is  no 
part  of  it  level  or  nearly  so.  It  is  almost  entirely  broken  by 
the  valleys  of  the  two  creeks  and  their  smaller  tributaries.  The 
height  above  the  sea  level  varies  from  620  feet  at  the  level  of 
the  Mississippi  river  in  the  southeastern  corner  of  Jefferson,  to 
1 200  feet  in  the  northwestern  corner  of  Mayville.  Crooked 
creek,  from  the  source  of  the  north  fork  to  its  discharge  into 
Bluff  slough,  is  about  eleven  miles  in  length.  It  drains  about 
65  square  miles  of  territory.  The  south  fork,  a  branch  about 
three  miles  long,  lies  entirely  in  Mayville.  Winnebago  creek 
from  the  Big  spring  near  its  source,  to  its  discharge  into  Min- 

353 


354  MINNESOTA    BOTANICAL    STUDIES. 

nesota  slough,  is  about  twelve  miles  in  length.  There  are 
three  small  branches,  one  of  which  has,  within  the  last  decade, 
become  considerably  smaller  than  it  formerly  was,  on  account 
of  the  drying  up  of  several  springs  near  its  source.  The 
amount  of  water  discharged  from  each  of  the  two  main  creeks 
during  the  summer  months  is  probably  not  less  than  1,500,000 
cubic  feet  per  day.  Neither  creek  is  very  susceptible  to 
changes  of  season,  but  either  one  will  rise  very  rapidly  after  a 
sudden  heavy  rain-fall  and  return  to  its  usual  level  in  a  few 
hours. 

The  bluffs  are  high  and  steep,  and  not  adapted  to  cultivation. 
(Plates  XXII.  and  XXVI.)  However,  many  of  the  ridges  are 
cultivated  and  form  some  of  the  best  farms  in  this  part  of  the  state. 
The  valleys  being  subject  to  overflow  and  the  bluffs  very  steep, 
by  no  means  the  entire  area  is  adapted  to  cultivation.  This 
condition  is  very  favorable  for  the  collection  of  native  plants. 

The  valleys  are  narrow,  in  no  place  exceeding  a  mile  in 
width  from  the  brow  of  one  bluff  to  the  brow  of  the  one  op- 
posite. 

In  taking  a  view  of  the  ecological  groups  of  the  plants  in- 
habiting this  region,  the  territory  may,  for  convenience,  be 
divided  into  river  valley,  creek  valleys  and  bluffs. 

The  river  valley  is  so  distinct  from  the  creek  valleys  that  it  is 
almost  imperative  that  it  be  considered  separately.  The  bluffs 
along  the  river  vary  somewhat  from  the  other  bluffs,  but  not 
sufficiently  to  warrant  a  division  into  river  bluff  and  creek 
bluff. 

River  valley. — In  the  river  valley  I  include  the  area  from  the 
foot  of  the  bluffs  on  one  side  of  the  river,  to  the  foot  of  the 
bluffs  on  the  opposite  side,  not  including  any  tributaries.  Along 
this  stretch  of  the  river,  from  New  Albin,  Iowa,  to  Brownsville, 
Minnesota,  the  valley  varies  from  three  to  five  miles  in  width. 
The  main  channel  of  the  river  is  from  one-half  a  mile  to  a  mile 
wide.  The  remainder  of  the  area  between  the  bluffs  is  formed 
of  islands,  sloughs  and  lakes  during  most  of  the  year.  (Plate 
XXV.,  B.)  During  the  spring  and  early  summer  the  whole  area 
is  generally  flooded  so  that  collection  can  be  carried  on  only  dur- 
ing the  late  summer  and  autumn.  The  river  channel  proper  is 
not  a  fruitful  field  for  the  collection  of  higher  plants.  The 
sluggish  sloughs,  lakes  and  ponds,  however,  offer  excellent 
conditions  for  such  collection. 


Wheeler :'  FLORA  OF  SOUTHEASTERN  MINNESOTA.  355 

For  consideration,  the  water  plants  of  the  river  valley  may  be 
classified  into  four  main  groups  :  plankton,  attached  submerged 
aquatic,  attached  aquatic  plants  with  natant  leaves,  and  adaptive 
shore  plants. 

PLANKTON. 

The  plants  forming  this  group  are  those  which  are  not  at- 
tached to  any  soil  substratum,  and  so  are  rarely  found  in  any  of 
the  swift-flowing  currents,  but  rather  on  the  surface  of  protected 
lakes  and  ponds  and  near  the  high  banks  of  sloughs,  where 
they  are  protected  from  rapid  currents  of  wind  and  water.  The 
plants  comprising  this  group  are  : 

Azolla  carolintana,  Sptrodcla  -polyrhiza, 

Ceratophyllum  demersum,  Utricularia  vulgaris. 

Lemna  minor , 

With  them  are  often  found  plants  of  Sagittaria  and  Pota- 
mogeton which  have  been  dislocated  from  their  original  position 
on  the  soil.  They  seem  to  grow  nearly  as  well  and  bloom 
nearly  as  profusely  as  when  attached.  In  this  condition  they 
form  part  of  the  plankton,  but  as  they  are  originally  attached 
and  ordinarily  remain  so,  I  have  not  included  them  in  the  list 
of  plankton  types. 

One  of  the  most  beautiful  and  interesting  plants  of  this  group 
is  the  small  heterosporous  fern,  Azolla  caroliniana.  In  the 
early  part  of  the  summer  it  is  green  or  but  slightly  red  in  color 
and  only  scattered  plants  or  very  small  patches  can  be  found. 
In  the  later  summer  and  autumn  it  covers  large  areas  of  water 
with  a  deep  red  pure  growth  or  mixed  with  the  duckweeds.  In 
restricted  areas  it  often  grows  so  rapidly  late  in  the  summer  that 
it  is  pushed  up  from  the  surface  of  the  water  and  forms  ridges 
and  bunches  above  the  water-level. 

ATTACHED  SUBMERGED  AQUATIC  PLANTS. 

The  floor  of  some  of  the  very  shallow  ponds  and  sloughs  is 
covered  with  a  growth  of  bassweeds  and  pondweeds  that  are  en- 
tirely immersed.  This  group  contains  but  few  species  of  the 
higher  plants.  The  species  collected  are  : 

Naias  flex  His )  Potamogeton  pusillus, 

Naias  g-uadalupensis,  Potamogeton  zosteraefolius. 


356  MINNESOTA    BOTANICAL    STUDIES. 

ATTACHED  AQUATIC  PLANTS  WITH  NAT ANT  LEAVES. 
Castalia  tuberosa,  Potamogeton  lonchttes, 

Nelumbo  hitea,  Potamogeton  natans, 

Nymphaea  advena,  Sagittaria  cuneata. 

Nearly  every  one  who  has  ever  visited  any  of  the  lakes  or 
rivers  of  Minnesota  is  acquainted  with  at  least  one  representa- 
tive of  this  group,  the  white  water-lily,  Castalia  tuberosa.  This 
with  the  Indian  lotus,  Nelumbo  lutea,  and  the  yellow  pond-lily, 
Nymphcea  advena,  all  of  which  are  members  of  the  water-lily 
family,  are  the  most  conspicuous  and  beautiful  of  our  river 
plants.  They  cover  large  areas  of  shallow  water  for  sometimes 
a  mile  or  more  in  extent.  It  may  be  of  interest  to  call  attention 
to  the  methods  of  adaptation  of  these  plants  to  their  aquatic 
habitat.  The  white  water-lily  and  the  yellow  pond-lily  carry 
their  natant  leaves  on  long  flexible  petioles  which  allow  the 
leaves  to  remain  upon  the  surface  for  variations  of  several  feet 
in  the  height  of  the  water.  The  Indian  lotus,  however,  carries 
the  leaf-blades  upon  stiff  strong  petioles  some  of  which  are  car- 
ried up  to  the  water  surface  and  others  are  raised  from  one  to 
three  feet  above  the  water.  (Plate  XXV.,  A.)  In  case  the  water 
rises  the  natant  leaves  are  destroyed  but  those  that  are  raised 
above  the  surface  remain  useful  to  the  plant  and  may  in  this  way 
be  caused  to  float.  The  projecting  leaves  are  not  conspicuously 
modified  in  any  way  from  those  that  were  originally  natant. 

Both  the  Indian  lotus  and  the  white  water-lily  are  abundant 
in  the  sloughs  of  the  Mississippi  river  at  Jefferson.  The  yellow 
pond-lily  is  not  so  abundant  as  either  of  the  other  two.  The 
Potamogetons  with  floating  leaves  may  be  found  growing  with 
the  water-lilies  or  in  small  patches  scattered  throughout  the 
sloughs.  They  never  cover  very  large  areas  to  the  exclusion 
of  other  plants. 

ADAPTIVE  SHORE  PLANTS. 

Alisma  -plantago-aquatica,  Sagittaria  latifolia, 

Eleocharis  acicularis,  Sagittaria  rigida, 

Nelumbo  lutea,  Scirpus  lacustris. 
Polygonum  emersum, 

The  plants  living  on  the  shores  of  the  lakes  and  sloughs  must 
adapt  themselves  to  life  under  the  varying  conditions  in  which 
they  may  be  placed  by  the  rise  and  fall  of  the  water.  During 


Wheeler :   FLORA  OF  SOUTHEASTERN  MINNESOTA.  357 

low  stages  they  may  be  left  out  of  the  water  entirely  and  when 
the  water  is  at  its  height  most  of  them  are  nearly  or  quite  sub- 
merged. The  plants  adapting  themselves  to  these  conditions 
might  be  considered  as  the  Sagittaria  group,  for  the  two  Sagit- 
tarias — latifolia  and  rigida — are  the  most  abundant  shore  plants 
with  the  possible  exception  of  Eleocharis  acicularis.  Nelumbo 
lutca  may  often  be  seen  in  times  of  very  low  water,  grow- 
ing on  the  muddy  banks  entirely  emersed  holding  its  leaves 
erect  two  or  three  feet  above  the  mud,  while  the  Castalia  when 
placed  under  these  conditions  lodges  its  leaves  on  the  mud  where 
they  soon  die.  Polygonum  emersum  covers  many  banks  to  the 
exclusion  of  other  vegetation.  It  is  adapted  to  living  on  the 
exposed  mud  or  in  the  water  but  under  the  latter  conditions  it 
always  projects  its  leaf-bearing  stems  out  of  the  water  and  keeps 
the  foliage  leaves  emersed. 

WET    MEADOWS    OF    THE    RIVER    VALLEY. 

During  a  large  part  of  the  growing  season  the  wet  meadows 
of  the  river  bottoms  are  submerged.  When  they  are  exposed 
for  a  sufficient  length  of  time  to  become  somewhat  dry  the 
grasses  are  generally  cut  for  hay.  The  plants  living  under 
these  conditions  are  mostly  coarse  grasses  and  sedges.  No  trees 
but  willows  seem  to  be  able  to  live  upon  these  meadows  and 
they  do  not  then  attain  tree  size.  Some  of  the  plants  forming 
the  vegetation  of  the  wet  meadows  are  : 

Asclepias  incarnata,  Scirpus  atrovirens^ 

Cyperus  esculentus,  Scirpus  cyperinus, 

Eleocharis  acicularis,  Stum  cicutaefolium, 

Elymus  virginicus,  Sparganium  eurycarpum, 

Eragrostis  hypnoides,  Spartina  cynosuroides, 

Eupatoriiim  purpureum ,  Vernonia  fasciculata , 

Homalocenchrus  virginicus,  Zizania  aquatica. 
Penthorum  sedoides, 

MUD-FLAT  VEGETATION. 

The  mud-flat  comprises  the  highest  land  of  the  islands.  It  is 
flooded  only  during  the  early  summer  but  on  account  of  its 
growth  of  timber  and  shrubs  the  soil  remains  wet  during  the 
entire  year.  The  largest  trees  growing  anywhere  in  this  region 
are  found  on  the  mud-flats  of  the  Mississippi  river. 


358  MINNESOTA    BOTANICAL    STUDIES. 

The  plants  which  form  large  trees  on  the  islands  are  : 

Acer  sacchartnum,  Populus  deltoides, 

Betula  nigra,  Qitcrcus  platanoides, 

Fraxinus  lanceolata,  Salix  amygdala  ides, 

Fraxinus  nigra,  Ulmus  americana. 
Gleditsia  triacanthos, 

The  following  species  do  not  attain  large  size,  but  are  either 
scattered  throughout  as  shrubs  or  small  trees,  or  form  a  dense 
low  growth  on  some  of  the  lower  grounds  of  the  mud  flat. 
(Plate  XXV.,  A.) 

Cephalantkus  occidentalism          Salix  Jluviatilis, 
Cornus  amonum,  Salix  nigra. 

Three  species  of  woody  vines  are  common  throughout  the 
islands.  The  Virginia  creeper,  Parthenocissus  quinquefolia, 
and  wild  grape,  Vitis  vulpina,  are  abundant,  covering  and  in 
many  cases  killing  large  trees.  The  climbing  poison  ivy,  Rhus 
radicans,  is  common  throughout  the  most  densely  wooded  parts. 
It  sometimes  climbs  to  a  height  of  twenty-five  or  thirty  feet,  and 
develops  a  stem  from  two  to  three  inches  in  diameter. 

During  the  late  summer  and  autumn  the  mud-flat  throughout 
is  covered  with  a  dense  growth  of  coarse  herbs  most  of  which 
are  perennials. 

The  following  herbaceous  plants  grow  on  the  mud-flat. 

Acnida  tamariscina,  Mimulus  ringens, 

Apocynum  cannabinum,  Onoclea  sensibilis, 

Ariscema  dracontium,  Polygonum  hartivrightii, 

Bidens  comosa,  Polygonum  hydropiper aides, 

Bidens  f rondo sa,  Polygonum  incarnatum, 

Bidens  Icevis,  Polygonum  punctatum^ 

Cicuta  bulbifera,  Polygonum  virginianum, 

Helenium  aulumnale,  Phy salts  philadelphica,, 

Ilysanthes  gratioloides,  Physostegia  virginiana, 

Lippia  lanceolata,  Ranunculus  pennsylvanicus , 

Lobelia  cardinalis,  Scutellaria  lateriflora, 

Lycopus  americanus,  Stachys  palustris, 

Lycopus  hicidus,  Steironema  ciliata, 

Lycopus  rubellus,  Teucrium  canadense, 

Lycopus  virginicus,  Urtica  gracilis, 

Ly thrum  alatuni,  Urticastrum  divaricatum. 
Mentha  canadensis, 


Wheeler :    FLORA  OF  SOUTHEASTERN  MINNESOTA.          359 

Creek  valleys. — The  valleys  of  the  creeks  present  an  entirely 
different  aspect  from  the  river  valley.  The  creeks  have  their 
own  well-defined  channels  to  which  they  hold  almost  the  year 
round.  High  waters  never  last  for  any  great  period  of  time. 
Those  which  are  caused  by  the  melting  of  the  snows  in  the 
spring  generally  last  from  about  noon  to  sun-down  while  those 
which  are  supplied  by  the  heavy  June  showers  generally  rise 
and  fall  during  the  night  or  very  early  morning.  The  damage 
done  to  vegetation  is  almost  restricted  to  the  floods  of  the  sum- 
mer months.  They  come  in  the  season  of  most  rapid  growth 
and  destroy  a  large  part  of  the  season's  growth  with  which  they 
come  in  contact.  The  areas  inundated  by  these  floods  are  never 
very  extensive  compared  to  those  along  the  river.  At  most 
points  along  the  valleys  the  gradual  rise  of  the  land  from  the 
creeks  to  the  bluffs  is  sufficient  to  prevent  the  formation  of 
ponds  and  lakes  by  the  rise  of  the  water.  The  alluvial  soils 
deposited  on  the  flats  do  not  dry  up  until  late  in  the  summer  and 
so  have  very  little  growth  besides  coarse  weeds.  They  are 
often  cultivated  but  there  is  always  the  danger  of  the  crops 
being  destroyed  by  high  water.  Most  of  the  best  cultivated 
fields,  in  the  valleys  are  on  the  table  lands  adjacent  to  the  foot 
of  the  bluffs.  They  are  generally  fertile,  are  protected  from 
high  water  and  hard  winds  and  are  not  in  a  position  to  wash  to 
any  great  extent.  The  steep  banks  on  the  north  edges  of  the 
table  lands  are  generally  wooded  and  bear  the  richest  and 
greatest  variety  of  plants  that  can  be  found  anywhere  in  this 
region.  The  table  lands  are  often  very  sharply  marked  off 
from  the  creek  bottoms  and  steep  bluffs.  Towards  the  heads  of 
the  creeks  the  table  lands  disappear  and  there  is  a  gradual  rise 
from  the  creeks  to  the  bluffs. 

The  water  vegetation  of  the  creek  valleys  is  almost  entirely 
limited  to  the  cold  water  plants  of  the  springs  and  small  streams. 
There  are  very  few  ponds  or  marshes  to  contain  still  water 
forms. 

The  vegetation  of  the  land  may  be  divided  into  that  of  the 
wet  meadow,  moist  woods  and  mesophytic  field.  The  wet  mea- 
dow is  about  on  a  level  with  the  banks  of  the  creeks.  It  never 
becomes  very  dry  and  on  the  lower  places  shows  some  of  the 
characters  of  a  marsh.  The  vegetation  of  the  moist  woods 
is  well  shown  on  the  wooded  banks  bordering  the  table  lands. 
Moist  woods  often  cover  some  of  the  protected  table  lands  and 


360  MINNESOTA    BOTANICAL    STUDIES. 

extend  for  some  distance  up  the  narrow  dark  ravines.  In  places 
where  timber  covers  the  flooded  areas  the  vegetation  is  similar 
to  that  of  the  mud  flat  on  the  islands  near  the  river.  The  vege- 
tation of  the  open  table  lands  I  have  called  mesophytic  field. 

COLD    SPRING    VEGETATION. 

The  valleys  of  Winnebago  and  Crooked  creeks  have  a  great 
many  springs  arising  from  the  bases  of  the  bluffs  throughout 
their  whole  length  but  perhaps  more  numerous  at  the  heads  of 
the  creeks  than  elsewhere.  Some  of  the  springs  that  outlet  in 
low  level  land  occasionally  form  small  cold  bogs  in  which  the 
ordinary  cold  water  plants  find  very  favorable  conditions  for 
growth.  A  large  spring  near  the  head  of  Clear  creek,  a  short 
branch  of  Crooked  creek  contains  the  greatest  abundance  of 
typical  cold  water  plants  of  any  spring  visited.  The  large 
creeks  do  not  contain  much  vegetation.  The  smaller  creeks 
often  contain  plants  similar  to  those  of  the  cold  springs. 

The  plants  characteristic  of  cold  running  water  are  : 


Batrachium  divaricattim,  Mimulus  jam 

Batrachium  trichophyllum,  Philotria  canadensis, 

Berula  erecta^  jRortpa  nasturtium, 

Cardamine  bulbosa,  Veronica  americana, 

Epilobiums  —  coloratum  and  adenocaulon  —  are  often  found 
growing  in  cold  spring  water  but  are  not  peculiar  to  this  local- 
ity as  they  are  also  found  growing  in  moist  soil.  None  of  the 
spring  plants  can  be  called  common  to  large  areas,  for  the  con- 
ditions necessary  for  their  growth  are  limited  in  extent. 

POND    VEGETATION. 

There  are  but  very  few  natural  ponds  along  the  creeks.  The 
ponds  are  generally  artificial  and  as  such  present  a  variety  of 
conditions  and  a  corresponding  variety  of  plants.  A  small 
natural  pond  in  a  bog  near  Crooked  creek  contains  all  it  can 
hold  of  the  yellow  pond-lily.  (Plate  XXVII.  ,  B.)  This  is  the 
only  place  in  which  any  of  the  water-lily  family  were  found 
outside  of  the  sloughs  and  lakes  of  the  Mississippi  river. 

WET  MEADOW  VEGETATION  OF  THE  CREEK  VALLEYS. 

The  wet  meadows  naturally  cover  a  very  large  part  of  the 
creek  valleys  but  under  present  conditions  most  of  them  are 


Wheeler:   FLORA  OF  SOUTHEASTERN  MINNESOTA.  361 

used  for  pasture,  or  where  they  can  be  easily  drained  for  culti- 
vation, though  they  are  of  course  in  constant  danger  of  being 
flooded.  Under  these  conditions  there  are  but  few  wet  meadows 
which  have  retained  their  original  vegetation.  Many  of  them 
under  continual  pasturing  have  grown  up  to  coarse  weeds  and 
grasses.  The  greatest  variety  of  plants  is  found  where  the  wet 
meadow  has  been  used  as  a  hay  meadow.  This  offers  more 
nearly  the  natural  conditions  for  such  plants  as  L ilium  cana- 
dense  (Plate  XXVII.,  A).  Habenaria  leucophcea,  Pediciilaris 
lanccolata,  Saxifraga  pennsylvanica,  Chelone  glabra,  Parnas- 
sia  caroliniana,  Onoclca  sensibilis  and  many  others  in  the  list. 
The  plants  which  grow  in  the  wet  meadows  are  : 

Angelica  atropurpurea,  Ly thrum  alatum, 

Aster  novce-anglicB,  Macrocalyx  nyctelea, 

Aster  prenanthoidcs,  Mimulus  ringens, 

Aster  puniceus,  Onoclea  sensibilis , 

Aster  sagittifolius,  Parnassia  caroliniana, 

Caltha  palustris,  Pedicularis  lanceolata, 
Cerastium  longipedunculatum,  Pimpinella  integerrima, 

Chelone  glabra,  Rudbeckia  laciniata, 

Cicuta  bulbifera,  Rudbeckia  triloba, 

Cicuta  maculata,  Rumex  acetosella, 

Doellingeria  umbellata,  Rumex  crispus, 

Dryopteris  thelypteris,  Saxifraga  pennsylvanica, 

Gentiana  crinita,  Silene  alba, 

Gentiana  jlavida,  Silphium  laciniatum, 

Habenaria  leucophcea,  Silphium  perfoliatum, 

Habenaria  psy codes,  Viola  obliqua, 

Lilitim  canadense,  Zizia  aurea. 
Lobelia  syphilitica, 

MOIST  WOODS  VEGETATION. 

As  previously  stated  the  most  typical  moist  woods  vegetation 
is  to  be  found  on  the  north  banks  of  the  table  lands.  The  tim- 
ber on  the  banks  has  much  of  it  been  left  uncut  and  offers  the 
very  best  conditions  for  the  survival  of  moist  woods  vegetation. 
In  the  list  of  moist  woods  plants  here  given  are  included  only 
those  collected  or  noted  from  a  single  location  in  Winnebago 
valley.  It  is  a  bank  about  one-half  a  mile  long  bordering  on 
the  table  land  for  the  greater  part  of  its  length.  Some  of  the 


362 


MINNESOTA    BOTANICAL    STUDIES. 


plants  listed  do  not  seem  to  be  typical  moist  woods  plants  and 
in  such  cases  they  have  probably  been  driven  to  the  margin  of 
the  thicket  by  the  cultivation  of  the  table  land  on  one  side  and 
by  the  high  water  of  the  creek  bottom  on  the  other.  The  plants 
of  the  moist  woods  on  this  bank  are  : 


Acer  negundo, 
Acer  nigrum, 
Actcea  alba, 
Actcea  r ubra, 
Adiantum  pedatum, 
A  dopogon  virgin  icum , 
Adoxa  moschatcllina, 
Agastache  scrophulari&folia, 
Agrimonia  hirsuta, 
Amelanchier  canadensis, 
Anemone  quinquefolia, 
Apios  apios, 

Apocynum  andros&mifolium, 
Aralia  nudicaulis, 
Aralia  racemosa, 
Ariscema  triphyllum, 
Asarum  canadense, 
Asclepias  ex  alt  at  a, 
Asclepias  incarnata, 
Asclepias  syriaca, 
Asplenium  acrostichoides, 
Aspleniumfilix-fcemina, 
Bicuculla  cucullaria, 
Bidens  frondosa, 
Bidens  comosa, 
Botrychium  Virginian  inn , 
Campanula  americana. 
Car  ex  rosea, 
Carpinus  caroliniana, 
Caulophyllum  thalictroides, 
Cerastium  longipedunculatum, 
Circcea  lutetiana, 
Clematis  virginiana, 
Cornus  candidissima, 
Cornus  rotundifolia, 


Cornus  stolonifcra, 
Corylus  americana, 
Crat&gus  ptinctata, 
Cratagus  tomentosa, 
Cypripedium  hirsutum , 
Cystopteris  bulbifera, 
Deringa  canadensis, 
Diervilla  diervilla, 
Epilobium  adenocaulon, 
Epilobium  color atum, 
Equisetum  arvense, 
Erigeron  pulchellus, 
Erythronimn  albidum, 
Euonymus  atropurpureus, 
Eupatorium  ageratoides, 
Falcata  comosa, 
Fragaria  americana, 
Galium  aparine, 
Galium  boreale, 
Galium  trifidum, 
Galium  triflorum, 
Geranium  maculatum, 
Geum  strictum, 
Habenaria  bracteata, 
Hepatica  a  cut  a, 
Heracleum  lanatum, 
Ifumulus  lupulus, 
Hydrophyllu  m  virgin  icum , 
Impatiens  aurea, 
Impatiens  biflora, 
Juglans  cinerea, 
Juglans  nigra, 
Lactuca  floridana, 
Lathyrus  ochroleucus, 
Lathyrus  venosus, 


Wheeler:    FLORA  OF  SOUTHEASTERN  MINNESOTA. 


363 


Lcptorchis  liliifolia, 
Lobelia  s vph ilitica , 
Loniccra  dioica, 
Loniccra  sullivantii, 
Mains  ioensis, 
Menispcrmum  canadensis, 
Mcntha  canadensis, 
Micrainpclis  lobata, 
MitcUa  diphylla, 
^.Yabciltis  albus, 
Nepcta  cat  aria, 
Onoclea  struthiopteris, 
Osmunda  claytoniana, 
Ostrya  virginiana, 
Ox alis  sir  id  a, 

Parth  en  o  ciss  us   qu  in  qu  efo  Ha , 
Pedicularis  canadensis, 
Peramium  pu&escens, 
Phlox  dh'aricata, 
Phrvnia  leptostachya, 
Podophyllum  peltatum, 
Pol  on  on  him  reptans, 
Polygonatum  commutatum , 
Polygonum  mcarnatum, 
Po  lygon  um  hy  dropipero  ides , 
Populus  grandidentata, 
Populus  tremuloides, 
Potentilla  canadensis, 
Primus  americana, 
Primus  nigra, 
Prunus  serotina, 
Prunella  vulgaris, 
Primus  virginiana, 
Pteris  aquilina, 
Pyrola  elliptica, 
Qiiercus  coccinea, 
Qtiercus  macro carpa, 

rubra, 

velutina, 


Ranunculus  abortivus, 
Ranunculus  septentrionalis, 
Rhus  glabra, 
Ribes  cynosbati, 
Ribcs  floridum , 
Ribes  iiva-crispa, 
Rubus  occidentalis, 
Rubus  villosus, 
Rudbeckia  laciniata, 
Rudbeckia  triloba, 
Salix  amygdaloides, 
Salix  jluv  iatilis , 
Sambucus  canadensis, 
Sanicrila  gregaria, 
Sanicula  marylandica, 
Silene  alba, 
Smilax  herbacea, 
Solidago  canadensis, 
Staphylea  trifolia, 
Syndesmon  thalictroides, 

Thalictrum  dioicum, 

Thalictrum  purpuresccns , 

Tilia  americana, 

Trillium  cernuum, 

Trillium  erectum, 
Triosteum  perfolialum, 

Ulmus  americana, 

Ulmus  fulva, 

Urtica  gracilis, 

Urticastrwn  divan' catum , 

Uvular ia  grandiflora, 

Vagnera  racemosa, 

Viburnum  lent  ago, 

Viola  pubescens, 

Viola  obliqua, 

Vitis  vulpina, 

Washingtonia  claytoni, 
Xanthoxylum  americana. 


364  MINNESOTA  BOTANICAL  STUDIES. 

MESOPHYTIC  FIELD   VEGETATION. 

The  mesophytic  field  vegetation  as  it  exists  in  the  creek  valleys 
to-day  is  almost  entirely  a  result  of  cultivation.  The  table  lands 
which  bear  the  plants  of  the  mesophytic  field  were  formerly  al- 
most entirely  wooded.  To-day  they  are  cleared  of  timber  and 
used  for  cultivation.  They  furnish  the  best  fields  for  cultivation 
in  the  whole  district.  They  are  not  subject  to  the  overflow  of 
the  bottom  lands,  nor  to  the  drought  of  the  ridges,  nor  to  the 
washouts  of  the  side-hills.  Being  so  extensively  cultivated  the 
plants  growing  upon  them,  which  are  not  themselves  cultivated, 
are  almost  confined  to  the  edges  of  fields  and  thickets.  Under 
such  conditions  a  list  of  plants  of  this  area  would  have  no  bear- 
ing upon  the  natural  ecological  groups. 

Bluffs. — The  bluffs  bordering  the  river  differ  from  those 
bordering  creek  valleys  in  being  steeper  and  in  having  many 
more  precipitous  cliffs.  The  brow  of  the  bluff  along  the  river 
for  almost  the  entire  distance  bordering  the  territory  covered  ex- 
cept where  interrupted  by  branch  valleys  or  ravines  is  one  al- 
most perpendicular  limestone  cliff,  varying  from  a  few  feet  to  a 
hundred  feet  in  height.  Cliffs  of  this  sort  are  not  so  common 
back  from  the  river.  The  vegetation  of  the  river  bluffs  differs 
to  some  extent  from  the  creek  bluffs  in  its  character.  Some 
of  the  common  forest  trees  of  the  lowland  of  the  creek  valleys, 
instead  of  growing  on  the  lowland  of  the  river  valley  inhabit 
the  foot  of  the  river  bluff.  The  proximity  of  the  river 
bluff  to  larger  areas  of  water  seems  to  raise  the  moisture 
content  of  the  soil  of  the  river  bluff  above  that  of  the  creek 
bluff  at  the  same  height  above  water  level.  The  growth 
then  of  such  a  tree  as  the  black  walnut  at  the  foot  of  the  river 
bluff  does  not  show  that  it  grows  here  under  more  arid  conditions 
than  in  the  creek  valley,  but  that  the  same  conditions  of  moisture 
in  the  soil  are  found  at  a  higher  level  on  the  river  bluff  than  on 
the  creek  bluff. 

On  all  bluffs  the  vegetation  shows  the  greatest  variation  with 
the  direction  of  the  slope.  Those  facing  from  south  to  west 
and  receiving  the  direct  rays  of  the  sun  from  noon  to  4  P.  M. 
are  generally  bare  of  trees  (Plate  XXII.,  B)  and  shrubs  while 
those  facing  from  north  to  east  are  generally  thickly  wooded 
(Plate  XXVI.,  B).  Ravines  with  their  greater  amount  of 
moisture  in  the  soil  and  greater  protection  from  winds  are 


Wheeler:   FLORA  OF  SOUTHEASTERN  MINNESOTA.  365 

generally  wooded  to  some  extent  whatever  the  direction  of  the 
slope  (Plate  XXII.,  A  and  B). 

Near  the  heads  of  the  creeks  at  the  bases  of  the  northern 
slopes  are  many  moist  limestone  cliffs  with  their  characteristic 
abundance  of  liverworts,  mosses  and  ferns,  sometimes  almost  to 
the  exclusion  of  the  higher  seed  plants.  The  moist  cliffs  bear 
more  of  the  northern  types  of  plants  rare  to  this  region  than 
any  one  other  special  area. 

The  zones  of  forest  vegetation  on  the  bluffs  are  often  very 
distinctly  marked  out  by  a  few  species.  The  oaks,  ^.  rubra, 
Q.  macrocarpa  and  <Q.  coccinea  extend  from  the  valley  to  the 
ridge  of  the  bluff  in  varying  degrees  of  abundance  and  thus 
do  not  determine  the  zone.  With  these,  however,  are  a  few 
species  which  are  limited  either  to  the  base  or  ridge.  The 
aspen  and  a  large-tooth  poplar  as  primary  and  the  ironwood, 
juneberry  and  wild  crab  as  secondary  species  mark  out  the  basal 
zone  of  forest  and  the  white  oak,  white  birch  and  shagbark 
hickory  in  varying  proportions  mark  out  the  ridge  forest.  Be- 
tween the  zone  containing  white  birch  on  the  ridge  and  that  con- 
taining the  aspen  at  the  base  is  a  zone  which  is  almost  entirely 
covered  with  dark-barked  trees.  Thus  the  zones  are  clearly 
shown  by  the  white  bark  of  the  white  birch  on  the  ridge  and 
that  of  the  aspen  below  with  the  dark-barked  trees  between. 

The  vegetation  areas  of  the  bluffs  may  be  considered  as  moist 
cliffs,  wooded  slopes  and  ravines,  ridge  forests,  bare  slopes  and 
open  ridges,  and  dry  rocks.  The  plants  of  the  moist  cliffs  are 
hydrophytic ;  those  of  the  wooded  slopes  and  ravines  which  in- 
clude a  large  part  of  the  bluff  area  are  mesophytic ;  the  ridge 
forests  are  xerophytic  and  the  bare  slope,  open  ridge  and  dry 
rock  plants  which  grow  on  the  most  exposed  and  dry  areas  in 
this  region  are  distinctly  xerophytic. 

MOIST    CLIFF    VEGETATION. 

This  group  of  plants  is  one  of  the  most  interesting  of  this  re- 
gion. One  is  always  on  the  lookout  for  rare  plants  to  this 'part 
of  the  state  from  the  secluded  and  often  almost  inaccessible 
moist  cliff.  The  short  list  of  plants  given  here  might  be  said 
to  be  almost  peculiar  to  moist  cliffs  as  they  are  rarely  found 
elsewhere.  Others  might  be  named  that  grow  upon  moist  cliffs, 
but  which  are  more  characteristic  of  moist  woods. 

Some  of  the  plants  characteristic  of  moist  cliffs  are  : 


366  MINNESOTA    BOTANICAL    STUDIES. 

Acer  spicatum,  Dirca  palustris, 

Adoxa  moschatcllina,  Pcllcea  stelleri, 

Betula  hit  Co.,  Viburnum  opulus. 
Circcea  alpina, 

VEGETATION  OF  WOODED  SLOPES  AND  RAVINES. 

This  group  of  plants  borders  and  perhaps  encroaches  upon 
the  moist  woods  of  the  valley  upon  the  one  hand  and  the  ridge 
forest  upon  the  other.  It  covers  a  large  area  but  does  not  repre- 
sent the  variety  of  species  that  are  found  in  the  moist  woods  of 
the  valley. 

Some  of  the  plants  of  the  wooded  slopes  and  ravines  are 

Asplenium  acrostichoides,  Lilium  timbellattim, 

Asplenium  jllix-focmina^  Lobelia  cordifolia, 

Castilleja  coccinea,  Lobelia  inflata, 

Cypripedium  hirsutum,  Onoclea  struthiopteris, 

Cypripedium  spectabilis,  Osmunda  claytoniana, 

Cystopteris  fragilis,  Panax  quinquefolia, 

Epilobitim  coloratum^  Pcdicularis  canadensis^ 

Erechtites  hieracifolia,  Polygonatum  commutatum , 

Eupatorium  ageratoides,  Pteris  aquilina, 

Falcata  comosa,  Rubus  occidentalis, 

Hieracium  umbellatum,  Rubus  vtllosus, 

Hieracium  scabrum,  Silene  stellata^ 

Hypericum  maculata,  Smilax  herbacea, 

Hypericum  majus,  Smilax  hispida. 

RIDGE  FOREST  VEGETATION. 

Most  of  the  woods  upon  the  ridges  are  rather  open  and  there- 
fore present  somewhat  xerophytic  conditions.  The  principal 
forest  trees  of  the  ridges  are  those  which  have  been  previously 
mentioned,  i.  e.,  Betula  papyrifera,  Hicoria  ovata,  Qitercus 
alba  and  JQuercus  macrocarpa.  Scattered  trees  of  other  kinds 
are  found  on  special  areas.  On  the  point  of  a  bluff  near  the 
village  of  Freeburg,  several  trees  of  Gymnocladus  dioicus  were 
found  in  one  of  the  most  exposed  locations  that  it  could  obtain. 
This  is  a  very  unusual  location  for  this  tree.  Prunus  virginiana, 
Juniperus  virginiana,  Tilia  americana  and  Celtis  occidentalis 
quite  frequently  grow  near  or  on  the  rocky  summits  of  the  bluffs 
but  do  not  cover  large  areas. 


Wheeler :   FLORA  OF  SOUTHEASTERN  MINNESOTA.          367 

BARE  SLOPE  AND  OPEN  RIDGE  VEGETATION. 

The  soil  of  the  southern  slope  and  open  ridge  is  generally 
largely  formed  of  sand  and  broken  limestone.  It  becomes  very 
dry  early  in  the  summer,  and  then  appears  almost  bare  of 
vegetation  except  where  it  is  broken  by  scattered  junipers 
(Plate  XXL,  A)  or  patches  of  Rhus  glabra. 

Some  of  the  plants  characteristic  of  the  bare  slope  and  open 
ridge  are  : 

Aceratcs  viridiflora,  Kuhnistera  purpurea, 

Asclepias  vcrticillata,  Lacinaria  cylindracea, 

Aster  sericeus,  Lacinaria  scarwsa, 

Boutcloua  curtipcndula,  Lappula  lappula, 

Boutcloua  hirsuta,  Linum  sulcatum, 

Coreopsis  palmata,  Lobelia  spicata, 

Cyperus  filiciilmis,  Oxalis  violacea, 

Cyperus  houghtoni,  Polygala  verticillata, 

Cyperus  schiveinitzii,  Polygonum  tenuc, 

Draba  caroliniana,  Piilsatilla  hirsutissima, 

Elymus  canadcnsis,  Ratibida  columnaris, 

Hclianthus  occidentalism  Rhus  glabra, 

Euphorbia  heterophylla,  Rhus  radicans, 

Hicracium  canadense,  Silene  antirrhina, 
Juniperus  commum's,  Valeriana  cdulis, 

Juniperus  sabina,  Viola  pedata, 

Kcelcria  cristata,  Viola  pedatifida. 

Kuhnistera  Candida, 

DRY  ROCK  VEGETATION  (Plate  XXL,  B). 

The  rock  plants  and  sand  plants  do  not  in  many  places  form 
distinct  groups.  The  sand  of  the  bluffs  nearly  always  contains 
considerable  broken  limestone  and  thus  furnishes  conditions 
favorable  to  the  growth  of  limestone  plants.  Pellcza  atropur- 
purea  and  Camptosorus  rhizophyllus  seem  to  be  the  only  ones 
that  are  restricted  to  the  bare  limestone.  The  former  prefers 
dryer  and  more  exposed  locations  than  the  latter. 

The  characteristic  plants  of  dry  rocks  are : 

Betula  papyri/era,  Juniperus  communis, 

Campanula  rotundifolia^  Pellaa  atropurpurea, 
Camptosorus  rhizophyllus^  Valeriana  edulis, 

Cystopteris  bulbifera,  Zygadenus  elegans. 


368  MINNESOTA    BOTANICAL    STUDIES. 

Results  of  the  survey. — The  botanical  survey  of  this  part  of 
the  state  was  undertaken  with  a  great  deal  of  interest  by  the 
collectors.  Never  before  has  this  region  been  explored  for  the 
purpose  of  botanical  collection.  Great  opportunities  were  there- 
fore offered  in  the  search  for  species,  which  may  have  made 
this  corner  of  the  state  the  northern  limit  of  their  range  and  for 
those  which  may  have  strayed  down  the  Mississippi  river  from 
their  native  home  at  its  headwaters.  With  such  possibilities  in 
view  the  collectors  were  not  disappointed  with  the  results. 

As  a  result  of  the  survey  578  species  of  plants  were  collected, 
26  of  which  are  Pteridophyta,  5  Archispermas,  87  Monocotyle- 
dons and  460  Dicotyledons. 

In  the  catalogue  of  species  are  reported  the  following  plants 
which  either  have  not  been  previously  reported  from  Minnesota 
or  have  been  reported  without  any  known  authentic  collection. 
The  specimens  have  been  placed  in  the  Herbarium  of  the  Uni- 
versity. 

Allionia  linear  is  ,\  Hicracium  umbellatumjt 

Asplenium  angustifolium,  Meibomia  illinoensis, 

J3idens  comosa,*  JVaias  guadalupensis, 

Car  ex  torta^  Primus  nigra, 

CratcBgus  macracantha,  J^tiercus  -prinoides, 

Falcata  pitcheri,  *  Rudbeckia  triloba, 

Gleditsia  triacanthos,  Sanicula  trifoliata, 

Helianthus  atrorubens^  Senecio  plattensis.* 

The  following  plants  collected  are  of  great  interest  as  rare 
plants  in  the  state  or  in  this  part  of  the  state. 

Arctostaphylos  uva-urst,  Gaura  biennis, 

Asclepias  obtusifolia,  Hamamelis  virginiana, 

Azolla  caroliniana,  Hydrocotyle  americana, 

Betula  lenta,  Jtmiperus  sabina, 

Carex  lurida^  Lactuca  ludoviciana, 

Cheilanthes  gracilis,  Lactuca  sagittifolia, 

Crat<zgus  punctata,  Meibomia  dillenii, 

Cyperus  houghtoni,  Polygonum  tenue, 

Dasystoma  grandijlora,  Polygonum  virginianum, 

Dryopteris  goldieana,  ^iiercus  platanoides, 

*  Previously  collected  but  not  reported  from  Minnesota. 

t  Previously  reported  from  incorrect  determinations  or  from  general  distri- 
bution ranges  given  in  large  manuals  without  authentic  collection. 


Wheeler  :   FLORA  OF  SOUTHEASTERN  MINNESOTA.  369 


Sanicula  gregaria,  Thalesia 

Sagittaria  cuneata,  Woods  ta  oregana. 

Soli  dago  crccta, 

CATALOG  OF  SPECIES  COLLECTED. 

The  following  catalog  of  plants  contains  only  those  collected  by 
Mr.  H.  L.  Lyon  and  the  writer  in  the  southeastern  part  of 
Houston  county.  With  the  exception  of  about  ten  species 
which  were  collected  in  Brownsville,  they  were  all  gathered  in 
the  townships  of  Mayville,  Crooked  Creek,  Winnebago  and 
Jefferson. 

The  determinations  were  almost  entirely  made  by  the  col- 
lectors, each  determining  the  plants  of  his  own  collection.  The 
determinations  of  the  species  of  Physalis  were  kindly  made  by 
Mr.  P.  A.  Rydberg,  of  Columbia  University,  and  those  of 
jQtiercus  prinoides,  velutina,  coccinea  and  rubra,  and  Betula 
lenta  by  Professor  C.  S.  Sargent. 

The  nomenclature  is  that  of  Britton  and  Brown's  Illustrated 
Flora  of  the  Northern  United  States  and  Canada. 


PTERIDOPHYTA. 

OPHIOGLOSSACEJE. 

Botrychium  virginianum  (L.)  Sw.    Schrad.   Journ.   Bot.  2: 

in.     1800. 

Coll.  :  Lyon  38,  Winnebago;  207,  Mayville.     June,  July. 
Infrequent,  rich  woods  and  shady  banks. 

OSMUNDACE^. 

Osmunda  claytoniana  L.  Sp.  PI.  1066.     1753. 

Coll.  :  Lyon  43,  Winnebago.     June. 
Common,  shady  hillsides  and  ravines. 

POLYPODIACE.E. 

Onoclea  sensibilis  L.  Sp.  PL  1062.     1753. 

Coll.  :  Lyon  326,  Jefferson.     Aug. 
Common,  wet  meadows  and  river  bottoms. 
Onoclea  struthiopteris  (L.)  HOFFM.  Deutsch.  Fl.  2  :  n.    1795. 
Coll.  :  Lyon  79,  Winnebago  ;  208,  Mayville.     June,  July. 
Common,  moist  thickets  and  river  bottoms. 


370  MINNESOTA    BOTANICAL    STUDIES. 

Woodsia  oregana  D.  C.  EATON,  Can.  Nat.  2 :  90.     1865. 

Coll.  :   Lyon  306,  Jefferson.     Aug. 

Rare  and  local,  on  brow  of  river  bluff.  The  only  previous 
collection  reported  from  Minnesota  is  that  from  Stillwater 
by  Miss  Field.  There  are  no  previously  collected  speci- 
mens from  Minnesota  in  the  Herbarium  of  the  University. 

Cystopteris  bulbifera  (L.)  BERNH.  Schrad.  Neues  Journ.  Bot. 

I  :  Part  2,  26.      1806. 
Coll.  :  Lyon  57,  Winnebago.     June. 
Common  on  shaded  rocks  and  limestone  ledges. 

Cystopteris  fragilis  (L.)  BERNH.  Schrad.  Neues  Journ.  Bot. 

I  :  Part  2,  27.      1806. 
Coll.  :  Lyon  221,  Mayville.     July. 
Frequent  in  deep  woods. 

Dryopteris  thelypteris  (L.)  A.  GRAY,  Man.  630.     1848. 

Coll.  :  Lyon  461,  Brownsville.     Aug. 
Common  in  swamps  and  wet  meadows  along  Wild  Cat  creek. 

Dryopteris  goldieana  (HOOK.)  A.  GRAY,  Man.  631.     1848. 

Coll.  :  Lyon  203,  222,  Mayville.     July. 

Rare  and  local,  deep  rich  woods.  The  only  previous 
authentic  collection  in  Minnesota  is  that  of  Leiberg  at  Min- 
neopa  falls,  Blue  Earth  County. 

Dryopteris  spinulosa  (RETZ.)   KUNTZE,   Rev.  Gen.  PL  813. 

1891. 

Coll.:  Lyon  253^,  Mayville.     July. 
Rare  and  local,  deep  rich  woods. 

Camptosorus  rhizophyllus   (L.)  LINK,   Hort.   Berol.   2:    69. 

1833- 

Coll.  :  Lyon  32,  65,  Winnebago.     June. 
Infrequent  or  rare,  limestone  ledges  and  boulders. 

Asplenium  angustifolium  MICHX.  Fl.  Bor.  Am.  2  :  265.     1803. 

Coll.  :  Lyon  204,  224,  Mayville.     July. 

Rare,  deep  rich  woods.  Not  previously  reported  from 
Minnesota. 

Asplenium   acrostichoides    Sw.    Schrad.   Journ.    Bot.    2 :   54. 

1800. 
Coll.:    Lyon   206,   223,  Mayville;    318,  Jefferson.     July 

Aug. 
Frequent,  rich  woods  and  moist  thickets. 


Wheeler:   FLORA  OF  SOUTHEASTERN  MINNESOTA.          371 

Asplenium  filix-fcemina  (L.)    BERNH.  Schrad.  Neues  Journ. 

Bot.  i :  Part  2,  26.     1806. 
Coll.  :   Lyon  205,  220,  Mayville.     July. 
Common  woods  and  thickets. 
Adiantum  pedatum  L.  Sp.  PI.  1095.     1753. 

Coll.  :  Lyon  45,  Winnebago.     June. 
Common,  woods  and  shady  banks. 
Pteris  aquilina  L.  Sp.  PI.  1075.     1753. 

Coll.  :  Lyon  42,  Winnebago.     June. 
Common,  hillsides  and  cut-over  timber  lands. 
Pellaea    stelleri    (S.    G.    GMEL.)   WATT,    Can.    Fil.    No.    2. 

1869-70. 

Coll.  :  Lyon  77,  Winnebago.     June. 
Infrequent,  moist  limestone  ledges. 
Pellaea  atropurpurea  (L.)   LINK,  Fil.  Hort.  Berol.  59.     1841. 

Coll.  :  Lyon  30,  Winnebago.     June. 
Frequent,  dry  limestone  cliffs  and  boulders. 
Cheilanthes  gracilis  (FEE)  METT.  Abh.  Senck.  Nat.  Gesell. 

3:    (reprint  36).     1859. 
Coll.  :  Lyon  299,  305,  Jefferson.     Aug. 
Rare  and  local,  dry  limestone  cliff.     There  is  no  previous 
authentic    collection  of    this  from  Minnesota  in  the  Uni- 
versity Herbarium.     Sandberg's  collection  from  Vermillion 
lake   made  in  1885  and  reported  as  this  species  should  be 
Cheilanthes  lanosa  (Michx.)  Watt,  which  has  not  previously 
been  reported  from  Minnesota. 
Polypodium  vulgare  L.  Sp.  PI.  1085.     1753. 

Coll.  :  Lyon  76,  Winnebago.     June. 
Local  on  limestone  ledge. 

SALVINIACE^:. 

Azolla  caroliniana  WILLD.  Sp.  PI.  5:  541.     1810. 

Coll.  :  Lyon  276,  Allamakee  Co.,  Iowa;   298,  444,  Jeffer- 
son.    Aug. 
Abundant  on  sloughs  and  lakes  of  the  Mississippi. 

EQUISETACE.E. 

Equisetum  arvense  L.  Sp.  PL  1061.     1753. 

Coll.  :  Lyon  102,  Winnebago.     June. 
Frequent,  meadows  and  pastures. 


372  MINNESOTA    BOTANICAL    STUDIES. 

Equisetum  pratense  EHRH.  Hanov.  Mag.  138.     1784. 

Coll.  :  Lyon  29,  Winnebago.     June. 
Frequent  in  light  shaded  soil. 

Equisetum  hyemale  L.  Sp.  PI.  1062.     1753. 

Coll.  :  Lyon  415,  Winnebago.     Aug. 
Common. 

Equisetum  laevigatum   A.  BR.  ;    Engelm.  Amer.  Journ.  Sci. 

46:  87.     1844. 

Coll.  :  Lyon  18,  Winnebago.     June. 
Local,  moist  meadows. 

SELAGINELLACEJE. 

Selaginella  rupestris  (L.)  SPRING,  in  Mart.  Fl.  Bras,  i :  Part 

2,  118.     1840. 

Coll.  :  Lyon  78,  Winnebago.     June. 
Infrequent,  dry  rocks. 


SPERMATOPHYTA. 

ARCHISPER1YLE. 

PINACE-3E. 

Pinus  strobus  L.  Sp.  PL  1001.     1753. 

Coll.  :  Wheeler  166,  254,  Winnebago.     June. 
Local  on  bluffs  along  Winnebago  and  Crooked  Creeks. 

Juniperus  communis  L.  Sp.  PI.  1040.     1753. 

Coll.:  Wheeler    108,    Winnebago;    213,  Jefferson ;    349, 

Crooked  Creek.     June,  July. 
Common  on  dry  bluffs.     (Plates  XXI. ,  A  and  B,  XXIV. ,  B.) 

Juniperus  virginiana  L.  Sp.  PI.  1039.     1753. 

Coll.  :  Wheeler  158,  169,  Winnebago.     June. 
Frequent  on  dry  bluffs. 

Juniperus  sabina  L.  Sp.  PL  1039.     I753- 

Coll.  :  Wheeler  214,  Jefferson.     June. 

Rare  and  local  on  dry  bluffs.  No  previous  collection  reported 
from  this  part  of  the  state.  This  is  about  the  most  southern 
point  of  collection  for  this  species  in  the  United  States  accord- 
ing to  Britton  and  Brown. 


Wheeler:   FLORA  OF  SOUTHEASTERN  MINNESOTA.          373 

TAXACE^E. 
Taxus  minor   (Micnx.)   BRITTON,  Mem.  Torr.  Club,  5 :    19. 

1893. 
Coll.  :  Wheeler  289,  Crooked  Creek;  317,  Mayville  ;  433, 

Jefferson.     July. 

Infrequent,  generally  on  dry  limestone  ridges,  occasionally 
in  woods.  Not  previously  reported  from  the  southern  part 
of  the  state. 

METASPERM^E. 

TYPHACE^E. 

Typha  latifolia  L.  Sp.  PI.  971.     1753. 

Coll.  :  Wheeler  266,  Winnebago.     June. 

SPARGANIACEJE. 

Sparganium  eurycarpum  ENGELM.  in  A.  Gray,  Man.  Ed.  2, 

430.     1856. 
Coll.  :  Wheeler  423,  Jefferson.     July. 

NAIADACEJE. 

Potamogeton  natans  L.  Sp.  PL  126.     1753. 

Coll.  :  Wheeler  460,  Jefferson.     Aug. 

Potamogeton  lonchites  TUCKERM.  Am.  Journ.  Sci  (II.)  6  :  226. 
1848. 

Coll. :  Wheeler  395,  488,  Jefferson.     Aug. 
Potamogeton  zosterafolius  SCHUM.  Enum.  PL  Saell.  50.     i8oi< 

Coll.  :  Wheeler  462,  490,  497,  Jefferson.     Aug. 
Potamogeton  pusillus  L.  Sp.  PL  127.     1753. 

Coll.  :  Wheeler  461,  Jefferson.     Aug. 
Naiasflexilis(WiLLD.)  ROST.  &  SCHMIDT,  Fl.  Sed.  384.     1824. 

Coll.  :  Lyon  329,  Jefferson.     Aug. 

Naias  guadalupensis  (SPRENG.)  MORONG,  Mem.  Torr.  Club,  3 : 
Part  2,  60.     1893. 

Coll.  :  Lyon  443,  Jefferson.     Aug. 
Not  previously  reported  from  Minnesota.     Sloughs  and  lakes 

of  the  Mississippi  river. 

ALISMACE^. 

Alisma  plantago-aquatica  L.  Sp.  PL  342.     1753. 
Coll.  :  Lyon  384,  Jefferson.     Aug. 


374  MINNESOTA    BOTANICAL    STUDIES. 

Sagittaria  latifolia  WILLD.  Sp.  PI.  4:  409.     1806. 

Coll.  :  Wheeler  123,  Winnebago ;  304. 
Crooked  Creek  ;  492,  Jefferson.     June-Aug. 
Sagittaria  cuneata  SHELDON,  Bull.  Torr.  Club,  20  :  283.  pi. 

159-     l893- 
Coll.:  Wheeler  491,  495,  Jefferson.     Aug. 

Not  previously  reported  from  this  part  of  the  state  or  the 

Mississippi  river.     Frequent  in  sloughs. 
Sagittaria  rigida PURSH,  Fl.  Am.  Sept.  397.     1814. 
Coll.  :  Wheeler  486,  Jefferson.     Aug. 

Sagittaria  cristata  ENGELM.  ;  Arthur,  Proc.  Davenport  Acad. 

4:  29.     1882. 
Coll.  :  Lyon  481,  Jefferson.     Aug. 

VALLISNERIACE^E. 

Philotria  canadensis  (Micnx.)  BRITTON,  Science  (II.)  2 :  5. 

1895. 
Coll.  :  Lyon  174,  Winnebago.     July. 

GRAMINE^E. 

Andropogon  furcatus  MUHL.  ;  Willd.  Sp.  PI.  4 :  919.     1806. 
Coll.  :  Wheeler  404,  Jefferson.     July. 

Chrysopogon  avenaceus    (Micnx.)    BENTH.  Journ.  Linn.  Soc. 

19:  73.     1881. 
Coll.  :  Lyon  354,  Jefferson.     Aug. 

Syntherisma  sanguinalis  (L.)  NASH,  Bull.  Torr.  Club,  22: 
420.     1895. 

Coll.  :  Wheeler  424,  Jefferson.     July. 
Panicum  crus-galli  L.  Sp.  PI.  56.     1753. 

Coll.  :  Wheeler  420,  Lyon  478,  Jefferson.     July,  Aug. 
Panicum  porterianum  NASH,  Bull.  Torr.  Club  22:  420.    1895. 

Coll.  :  Wheeler  388,  Jefferson.     July. 

Panicum  scribnerianum  NASH,  Bull.    Torr.    Club,    22:    421. 
1895. 

Coll.  :  Wheeler  182,  Winnebago.     June. 
Panicum  virgatum  L.  Sp.  PL  59.     1753. 

Coll.  :  Wheeler  421,  Jefferson.     July. 
Panicum  capillare  L.  Sp.  PL  58.     1753. 

Coll.:  Lyon  477,  Jefferson.     Aug. 


Wheeler:   FLORA  OF  SOUTHEASTERN  MINNESOTA.          375 

Cenchrus  tribuloides  L.  Sp.  PI.  1050.     1753. 

Coll.  :  Lyon  287,  Jefferson.     July. 
Zizania  aquatica  L.  Sp.  PI.  991.     1753. 

Coll.  :  Wheeler  523,  Jefferson.     Aug. 

Homalocenchrus  virginicus  (WILLD.)  BRITTON,  Trans.  N.  Y. 
Acad.  Sci.  9:   14.      1889. 

Coll.  :  Wheeler  564,  Jefferson.     Aug. 
Spartina  cynosuroides  (L.)  WILLD.  Enum.  80.     1809. 

Coll.  :  Wheeler  426,  Jefferson.     July. 

Bouteloua  hirsuta  LAG.  Var.   Cienc.  y  Litter  2:  Part  4,  141. 
1805. 

Coll.  :  Wheeler  347,  Crooked  Creek;  Lyon  291, Jefferson. 

July- 

Bouteloua  curtipendula  (MICHX.)  TORR.   Emory's  Rep.  153. 
1848. 

Coll. :  Wheeler  362,  Crooked  Creek.     July. 
Eragrostis   hypnoides  (LAM.)  B.S.P.   Prel.   Cat.    N.  Y.  69. 
1888. 

Coll.  :  Wheeler  524,  Jefferson.     Aug. 
Koeleria  cristata  (L.)  PERS.  Syn.  i :  97.     1805. 

Coll.  :  Lyon  113,  Winnebago.     June. 
Panicularia  americana  (TORR.)  MACM.  Met.  Minn.  81.    1892. 

Coll.  :  Lyon  59,  Winnebago.     June. 
Bromus  ciliatus  L.  Sp.  PL  76.     1753. 

Coll.  :  Lyon  414,  Winnebago.     Aug. 
Bromus  kalmii  A.  GRAY,  Man.  600.     1848. 

Coll.  :  Lyon  259,  Jefferson.     July. 
Bromus  secalinus  L.  Sp.  PI.  76.     1753. 

Coll.  :  Lyon  184,  Winnebago.     July. 
Elymus  virginicus  L.  Sp.  PI.  84.     1753. 

Coll.  :  Wheeler  418,  Jefferson.     July. 
Elymus  canadensis  L.  Sp.  PI.  83.     1753. 

Coll.  :  Wheeler  292,  Mayville.     July. 

CYPERACE^. 

Cyperus  schweinitzii  TORR.  Ann.  Lye.  N.  Y.  3 :  276.     1836. 

Coll.  :  Lyon  375,  Jefferson.     Aug. 
Cyperus  esculentus  L.  Sp.  PI.  45.     1753. 

Coll.  :  Wheeler  526,  Jefferson.     Aug. 


376  MINNESOTA    BOTANICAL    STUDIES. 

Cyperus  filiculmis  VAHL,  Enum.  2  :  328.     1806. 
Coll.  :  Wheeler  348,  Crooked  Creek.     July. 

Cyperus  houghtoni  TORR.  Ann.  Lye.' N.  Y.  3:   277.     1836. 

Coll.  :  Wheeler  346,  Crooked  Creek.     July. 
The  only  previous  collection  known  from  Minnesota  is  that 
of  Holzinger,  St.  Croix  River,  Minn.     Britton  reports  this 
collection  in  the  Bull.  Torr.  Club,  18 :  368.      1891. 
The  collection  from  Crooked  Creek  was  made  from  the  sum- 
mit of  a  very  dry  sandy  hill.     Both    C.  houghtoni  and  C. 
schiveinitzii  grow  in  sand  but   the  former  probably  grows 
in  the  drier  locality  of  the  two. 

Eleocharis  acicularis  (L.)  R.  &  S.  Syst.  2  :  154.     1817. 
Coll.  :  Wheeler  527,  Jefferson.     Aug. 

Scirpus  lacustris  L.  Sp.  PI.  48.     1753. 

Coll.  :  Wheeler  148,  Winnebago.     June. 

Scirpus  atrovirens  MUHL.  Gram.  43.     1817. 
Coll.  :  Wheeler  267,  Winnebago.     June. 

Scirpus  cyperinus  (L.)  KUNTH,  Enum.  2:  170.     1837. 

Coll.  :  Lyon  168,  Crooked  Creek  ;  Wheeler  425,  Jefferson. 
June,  July. 

Carex  lupulina  MUHL.  ;   Schk.   Riedg.  2  :  54.  f.  123.     1806. 
Coll.  :  Lyon  280,  Jefferson.     July. 

£arex   lurida  WAHL.   Kongl.    Acad.    Handl.   (II.)   24:    153. 

1803. 

Coll.  :  Wheeler  142,  Winnebago.     June. 
No  Minnesota  specimens  in  the  Herbarium  of  the  University. 
Previously    collected  at  Lake  Itasca,  Sandberg  No.  1180. 

Carex  hystricina  MUHL.  ;  Willd.  Sp.  PI.  4:  282.     1805. 
Coll.  :  Wheeler  119,  Winnebago.     June. 

Carex filiformis  L.  Sp.  PL  976.     1753. 

Coll.  :  Wheeler  121,  Winnebago.     June. 

Carex  torta  BOOTT  ;  Tuckerm.  Enum.  Meth.  n.     1843. 

Coll.  :  Lyon  60,  Winnebago.     June. 

Not  previously  reported  from  Minnesota.  The  nearest  point 
of  previous  collection,  as  shown  by  the  Herbarium  of 
the  University,  is  Winnebago  county,  Wisconsin. 

Carex  stipata  MUHL.  ;   Willd.  Sp.  PL  4:  233.     1805. 
Coll.:  Wheeler  116,  Winnebago.     June. 


Wheeler:   FLORA  OF  SOUTHEASTERN  MINNESOTA.          377 

Carex  vulpinoidea  MICIIX.  Fl.  Bor.  Am.  2:  169.     1803. 

Coll.  :  Wheeler  144,  Winnebago.     June. 
Carex  rosea  Schk.  Riedgr.  Nachtr.  15.  f.  ijg.     1806. 

Coll.  :   Wheeler  n,  143,  Winnebago.     June. 
Carex  tribuloides  WAHL.  Kongl.  Vet.  Acad.  Handl.  (II.)  24: 
145.     1803. 

Coll.  :  Wheeler  118,  Winnebago.     June. 
Carex  cristatella  BRITTON,  Br.  &  Br.  111.  Fl.  N.  U.  S.  &  Can. 

I  :  357-  /•  865.      1896. 

Coll.  :  Wheeler  197,  Winnebago.     June. 

ARACE.E. 
Arisaema  triphyllum  (L.)  TORR.  Fl.  N.  Y.  2  :  239.     1843. 

Coll.  :  Lyon  31,  Winnebago.     June. 
Arisaema  dracontium  (L.)  SCHOTT,  Melet.  i :   17.     1832. 

Coll.  :  Lyon  239,  Mayville;   248.     Crooked  Creek.     July. 
The  only  previous  collections  reported  from  Minnesota  are 
Manning,  Lake  Pepin  and  Holzinger,  Winona.     Frequent 
in  moist  woods  along  the  Mississippi  River. 

LEMNACE.E. 
Spirodela  polyrhiza  (L.)  SCHLEID.     Linnaea,  13:   392.      1839. 

Coll.  :  Wheeler  113,  Winnebago.     June. 
Lemna  minor  L.  Sp.  PL  970.     1753. 

Coll.  :  Wheeler  112,  Winnebago.     June. 

COMMELINACE^). 
Tradescantia  virginiana  L.  Sp.  PI.  288.     1753. 

Coll.  :  Lyon  45,  Winnebago.     June. 
Tradescantia  reflexa  Raf.  Atl.  Journ.  150.     1832. 

Coll.  :  Wheeler  410,  Jefferson.     July. 
Not  previously  reported  from  Minnesota. 

JUNCACEJE. 

Juncus  effusus  L.  Sp.  PI.  326.     1753. 

Coll.  :  Lyon  58,  Winnebago.     June. 
Juncus  tenuis  WILLD.  Sp.  PI.  2:  214.     1799- 

Coll.  :  Lyon  73,  Winnebago.     June. 

MELANTHACE.E. 

Zygadenus  elegans  PURSH,  Fl.  Am.  Sept.  241.     1814. 
Coll.  :   Lyon  49,  Winnebago.     June. 


378  MINNESOTA    BOTANICAL    STUDIES. 

Uvularia   grandiflora  J.  E.  SMITH,  Ex.  Bot.  i  :    99.  pi.  51. 

1804-5. 
Coll.  :  Lyon  91,  Winnebago.     June. 


Lilium  umbellatum  PURSH,  Fl.  Am.  Sept.  229.     1814. 

Coll.  :  Lyon  146,  Winnebago.     June. 

All  previous  collections  of  this  species  from  Minnesota  have 
been  reported  as  L.  -philadelphicum  L.  The  latter  species 
so  far  as  known  has  not  been  collected  in  Minnesota. 

Lilium  canadense  L.  Sp.  PI.  303.     1753. 

Coll.  :  Lyon  199,  Crooked  Creek.     July. 
Common,  moist  meadows.     (Plate  XXVII.,  A.) 

Erythronium  albidum  NUTT.  Gen.  i  :  223.     1818. 
Coll.  :   Herb.  Wheeler  i,  Winnebago.     May. 

CONVALLARIACE^:. 

Vagnera  racemosa  (L.)  MORONG,  Mem.  Torr.  Club,  5:  114. 

1894. 
Coll.  :  Wheeler  67,  Winnebago.     June. 

Vagnera  stellata  (L.)  MORONG,  Mem.  Torr.  Club,  5:    114. 

1894. 
Coll.  :  Wheeler  97,  Winnebago.     June. 

Unifolium  canadense  (DESF.)  GREENE,  Bull.  Torr.  Club,  15  : 

287.     1888. 
Coll.  :  Wheeler  98,  Winnebago.     June. 

Polygonatum  commutatum  (R.  &  S.)  DIETR.  ;  Otto  &  Dietr. 

Gartenz.  3  :   222.     1835. 

Coll.:    Wheeler    78,    184,   Winnebago;    570,   Jefferson; 
Lyon  166,  Crooked  Creek.     June,  Aug. 

Trillium  erectum  L.  Sp.  PL  340.     1753. 
Coll.  :  Lyon  17,  Winnebago.     June. 

Trillium  cernuum  L.  Sp.  PL  339.     1753. 

Coll.  :   Herb.  Wheeler  2,  Winnebago.     May. 

SMILACE^E. 

Smilax  herbacea  L.  Sp.  PL  1030.     1753. 

Coll.:    Wheeler   376,    Mayville  ;     467,   Jefferson.     July, 
Aug. 


Wheeler:   FLORA  OF  SOUTHEASTERN  MINNESOTA.  379 

Smilax  hispida  MUHL.  ;  Torr.  Fl.  N.  Y.  2:  302.     1843. 

Coll.:   Wheeler  372,  Crooked   Creek;    Lyon  258,  Jeffer- 
son.    July. 

AMARYLLIDACE^. 
Hypoxis  hirsuta  (L.)  COVILLE,  Mem.  Torr.  Club,  5:     118. 

1894. 
Coll.  :  Wheeler  91,  Winnebago.     June. 

DIOSCOREACEJE. 

Dioscorea  villosa  L.  Sp.  PI.  1033.     1753. 

Coll.:    Wheeler    322,    May  ville ;    364,   Crooked    Creek. 

July- 

IRIDACEJE. 
Iris  versicolor  L.  Sp.  PL  39.     1753. 

Coll.  :  Lyon  362,  Jefferson.     Aug. 
Sisyrinchlum  angustifolium  MILL.  Gard.  Diet.  Ed.  7.      1759- 

kColl.  :  Lyon  75,  Winnebago.     June. 
ORCHIDACEJE. 
Cypripedium  reginae  WALT.  Fl.  Car.  222.     1788. 

Coll.  :  Wheeler  192,  Winnebago.     June. 
Cypripedium  candidum  WILLD.  Sp.  PL  4:  142.     1805. 

Coll.  :  Wheeler  99,  Winnebago.     June. 
Cypripedium  hirsutum   MILL.    Gard.    Diet.    Ed.    8,    No.    3. 

1768. 

Coll.  :  Wheeler  66,  82,  Winnebago.     June. 
Orchis  spectabilis  L.  Sp.  PL  943.     1753. 

Coll.  :   Herb.  Wheeler  8,  Winnebago.     June. 
Habenaria  bracteata  (WILLD.)  R.  Br.  in  Ait.  Hort.  Kew,  Ed. 

2,  5:    192.      1813. 

Coll.  :  Wheeler  106,  Winnebago.     June. 
Habenaria    leucophaea  (NUTT.)  A.  GRAY,  Man.   Ed.  5,  502. 

1867. 

Coll.  :  Wheeler  299,  Crooked  Creek.     July. 
Habenaria  psycodes  (L.)  A.  GRAY,  Am.  Journ.  Sci.  38:  310. 

1840. 

Coll.:  Wheeler  386,  Jefferson.     July. 

Peramium  pubescens  (WILLD.)  MAcM.  Met.  Minn.  172.    1892. 
Coll.  :  Lyon  100,  Winnebago.     June. 


380  MINNESOTA    BOTANICAL    STUDIES. 

Leptorchis  liliifolia  (L.)  KUNTZE,  Rev.  Gen.  PL  671.     1891. 
Coll.  :  Wheeler  107,  168,  195,  Winnebago ;  350,  Crooked 
Creek;  391,  Jefferson.     June,  July. 

JUGLANDACE^E. 

Juglans  nigra  L.  Sp.  PL  997.     1753. 

Coll.  :  Lyon  243.  Crooked  Creek.     July. 

Juglans  cinerea  L.  Sp.  PL  Ed.  2,  1415.     1763. 
Coll.  :  Lyon  62,  108,  Winnebago.     June. 

Hicoria  minima  (MARSH.)  BRITTON,  Bull.  Torr.  Club,  15  :  284. 

1888. 

Coll.:   Lyon   149,  Winnebago ;   238,  239,  Mayville  ;   475, 
Jefferson.     June,  Aug. 

Hicoria  ovata  (MILL)  BRITTON,  Bull.    Torr.  Club,   15:    283. 

1888. 
Coll.  :  Lyon  71,  Winnebago;  474,  Jefferson.     June,  Aug. 

SALICACE.E. 

Populus  alba  L.  Sp.  PL  1034.     1753. 

Coll.  :  Lyon  159,  Winnebago.     June. 

Populus  balsamifera  candicans  (AiT.)  A.  GRAY,  Man.  Ed.  2, 

419.     1856. 
Coll.  :  Lyon  156,  Winnebago.     June. 

Populus  grandidentata  MICHX.  Fl.  Bor.  Am.  2:  243.    1803. 
Coll.  :  Lyon  64,  Winnebago.     June. 

Populus  tremuloides  MICHX.  FL  Bor.  Am.  2:  243.     1803. 
Coll.  :  Lyon  88,  Winnebago.     June. 

Populus  deltoides  MARSH,  Arb.  Am.  106.     1785. 
Coll.  :  Lyon  125,  Winnebago.     June. 

Salix  nigra  MARSH,  Arb.  Am.  139.     1785. 
Coll.  :  Wheeler  265,  Winnebago.     June. 

Salix  amygdaloides  ANDERS.  Ofv.   Handl.  Vet.  Akad.  1858 : 

114.     1858. 
Coll.  :  Wheeler  137,  Winnebago.     June. 

Salix  lucida  MUHL.  Neue  Scrift.  Ges.  Nat.  Fr.  Berlin,  4:   239. 

fl.6.f.?.     1803. 

Coll.:  Wheeler  138^-,  Winnebago ;   232,  Crooked  Creek. 
June,  July. 


Wheeler:   FLORA  OF  SOUTHEASTERN  MINNESOTA.  381 

Salix  fluviatilis  NUTT.  Sylva,  i :  73.     1842. 

Coll.:   Wheeler  136,  Winnebago ;    333,  Crooked  Creek. 
June,  July. 

Salix  bebbiana  SARG.  Gard.  &  For.  8  :  463.     1895. 
Coll.  :  Wheeler  343,  Crooked  Creek.     July. 

Salix  humilis  MARSH,  Arb.  Am.  140.     1785. 
Coll.  :  Wheeler  181,  Winnebago.     June. 

Salix  discolor  MUHL.  Neue  Schrift.  Ges.  Nat.  Fr.  Berlin,  4 : 

234.^7.  6.  f.  i.     1803. 

Coll.:  Wheeler   138,  Winnebago;    334,  Crooked  Creek. 
June,  July. 

BETULACE^E. 

Carpinus  caroliniana  WALT.  Fl.  Car.  236.     1788. 
Coll.  :  Lyon  56,  Winnebago.     June. 

Ostrya  virginiana  (MILL.)  WILLD.  Sp.  PL  4:  469.     1805. 
Coll.  :  Wheeler  200,  Winnebago.     June. 

Corylus  americana  WALT.  Fl.  Car.  236.     1788. 
Coll.  :  Wheeler  22,  Winnebago.     June. 

Corylus  rostrata  AIT.  Hort.  Kew.  3:  364.     1789. 

kColl.  :    Wheeler  223,  Winnebago ;    275,  Crooked  Creek. 
June. 
Not  previously  collected  from  southern  part  of  state.     Local 
on  bluffs. 

Betula  papyrifera  MARSH.  Art.  Am.  19.     1785. 

Coll.:  Wheeler  215,  Jefferson.     June. 
Common,  dry  ridges.     (Plates  XXIII.,  A  and  XXIV.,  B.) 

Betula  nigra  L.  Sp.  PI.  982.     1753. 

Coll.  :  Wheeler  553,  Jefferson.     Aug. 
Common  in  the  lowlands  of  the  Mississippi  River. 

Betula  lenta  L.  Sp.  PI.  983.     1753. 

Coll.  :  Wheeler  165,  Winnebago.     June. 
Not  previously  collected  in  the  southern   part  of  the  state. 
Rare. 

Betula  lutea  MICHX.  f.  Arb.  Am.  2  :   152.  pL  5.     1812. 

Coll.:  Wheeler  199,  Winnebago;   271,  Crooked    Creek; 

325,  Mayville.     June,  July. 

Frequent  in  moist  locations  along  Winnebago  and  Crooked 
creeks. 


382  MINNESOTA    BOTANICAL    STUDIES. 

Betula  pumila  L.  Mant.  124.     1767. 

Coll.  ;  Wheeler  272,  Crooked  Creek.     June. 
Local  along  Crooked  creek,   forming  large  thickets  in  wet 

meadows. 
Alnus  incana  (L.)  WILLD.  Sp.  PL  4:  335.     1805. 

Coll.  :  Wheeler  617,  Brownsville.     Aug. 
Local  at  mouth  of  Wild  Cat  creek. 

FAGACEM. 
Quercus  rubra  L.  Sp.  PL  996.     1753. 

Coll.  :  Wheeler  640,  641,  Jefferson.     Aug. 
Common  throughout. 
Quercus  coccinea  WANG.  Amer.  44.  -pi.  ^..f.  p.     1787. 

Coll.  :  Wheeler  644,  645,  Jefferson.     Aug. 
Common  throughout. 

Quercus  velutina  LAM.  Encycl.  i:  721.  1783. 
Coll.  :  Wheeler  642,  643,  Jefferson.  Aug. 
Prof.  Sargent  writes  about  643  :  "  Collection  643,  which  I 
call  Q.  veluttna,  differs  from  that  species  as  it  usually 
occurs  by  the  much  smaller  less  tomentose  buds ;  the 
acorns,  however,  are  clearly  from  Q.  velutina.  I  fre- 
quently have  seen  specimens  of  this  same  form  from  the 
region  immediately  west  of  the  Great  Lakes.  It  appears 
sometimes  as  if  it  might  be  a  hybrid  between  Q  velutina 
and  J^.  coccinea  but  its  occurrence  is  too  frequent  and  its 
distribution  too  wide  to  admit  of  this  supposition.  With 
the  present  state  of  our  knowledge  I  can  but  refer  it  to  JQ. 
velutina.'''' 

Q.  velutina  does   not  seem  to  be   nearly  so   common  in  this 
region  as  Q  coccinea. 

Quercus  alba  L.  Sp.  PL  996.     1753. 

Coll.  :  Wheeler  638,  Jefferson.     Aug. 
Common  on  ridges  of  bluffs  throughout. 

Quercus  macrocarpa  MICHX.  Hist.  Chen.  Am.  2.  $1.  23.     1801. 

Coll.  :  Wheeler  639,  Jefferson.     Aug. 
Common  throughout. 

Quercus  platanoides  (LAM.)  SUDW.  Rep.   Secy.  Agric.   1892  : 

327.     1893. 

Coll.:  Wheeler    366,    Crooked    Creek;    456,    538,    654, 
Jefferson.     July,  Aug. 


Wheeler:   FLORA  OF  SOUTHEASTERN  MINNESOTA.  383 

No  previous  collection  reported  from  Minnesota.  Reported 
by  Garrison  as  frequent  at  several  points  near  the  head- 
waters of  the  Mississippi.  Frequent  on  the  lowlands  of 
the  Mississippi  in  Jefferson  and  Crooked  Creek  townships 
and  in  Allamakee  Co.,  Iowa. 

Quercus  prinoides  WILLD.  Neue  Schrift.  Ges.  Nat.  Fr.  Berlin, 

3:  397.     1801. 

Coll.  :  Wheeler  360,  Crooked  Creek.     July. 
Not  previously  reported  from   Minnesota.     Whether  this  is 
the   species  reported  by  Lapham   as  Jj>.   castanea  Willd. 
cannot  be  ascertained  as  there  are  no  specimens  from  Lap- 
ham's  collection  in  the  Herbarium  of  the  University. 
The  specimens  were  collected  from  two  trees  on  the  side  of  a 
bluff  in  section  19,  township  102  N.,  range  4  W. 

ULMACE^E. 

Ulmus  americana  L.  Sp.  PI.  226.     1753. 

Coll.  :  Wheeler  24,  Winnebago.     June. 

Ulmus  racemosa  THOMAS,  Am.  Jour.  Sci.  19  :  170.     1831. 

Coll.  :  Wheeler  315,  Mayville.     July. 
Infrequent  on  lowland  near  Crooked  creek. 

Ulmus  fulva  MICHX.  Fl.  Bor.  Am.  i :    172.     1803. 
Coll.  :   Wheeler  23,  Winnebago.     June. 

Celtis  occidentalis  L.  Sp.  PL  1044.     1753. 

Coll.  :  Wheeler   240,  Winnebago ;   278,   Crooked  Creek ; 
Lyon  374,  Jefferson.     June,  Aug. 

MORACE^E. 

Morus  rubra  L.  Sp.  PI.  986.     1753. 

Coll.  :   Lyon  368,  Jefferson.     Aug. 
Infrequent  along  Mississippi  river. 

Humulus  lupulus  L.  Sp.  PI.  1028.     1753. 
Coll.  :   Lyon  312,  Winnebago.     Aug. 

Cannabis  saliva  L.  Sp.  PI.  1027.     1753. 
Coll.  :  Lyon  282,  Jefferson.     Aug. 

URTICACE^E. 

Urtica  gracilis  AIT.  Hort.  Kew.  3:  341.     1789. 
Coll.  :  Lyon  126,  Winnebago.     June. 


384  MINNESOTA    BOTAXICAL    STUDIES. 

Urticastrum   divaricatum   (L.)   KUNTZE,  Rev.  Gen.  PI.  635. 

1891. 

Coll.:    Lyon    117,    Winnebago ;    358,   Jefferson.      June, 
Aug. 

Adicea  pumila  (L.)  RAF.  ;  Torr.   Fl.  N.  Y.  2  :   223.     As  syn- 
onym.     1843. 

Coll.:    Wheeler    327,    Mayville ;    653,   Jefferson.     July, 
Aug. 

Parietaria   pennsylvanica   MUHL.  ;    Willd.    Sp.    PI.   4;  955. 

1806. 
Coll.  :  Lyon  191,  Crooked  Creek.     July. 

SANTALACE^E. 

Comandra  umbellata  (L.)  NUTT.  Gen.  i:  157.     1818. 
Coll.  :  Lyon  90,  Winnebago.     June. 

ARISTOLOCHIACEJE. 

Asarum  canadense  L.  Sp.  PI.  442.     1753. 
Coll.  :  Wheeler  57,  Winnebago.     June. 

POLYGONACE^E. 

Rumex  acetosella  L.  Sp.  PI.  338.     1753. 
Coll.  :  Lyon  130,  Winnebago.     June. 

Rumex  crispus  L.  Sp.  PL  335.     1753. 

Coll.  :  Lyon  127,  158,  Winnebago.     June. 

Polygonum  hartwrightii  A.  GRAY,  Proc.  Am.  Acad,  8 :  294. 

1870. 
Coll.  :  Wheeler  606,  Brownsville.     Aug. 

Polygonum  emersum  (Micnx.)  BRITTON,  Trans.  N.  Y.  Acad. 

Sci.  8:  73.     1879. 
Coll.  :  Wheeler  394,  458,  Jefferson.     July,  Aug. 

Polygonum  incarnatum  ELL.  Bot.  S.  C.  &  Ga.  i :  456.     1817. 
Coll.  :   Wheeler  419,  Jefferson.     July. 

Polyonum  hydropiper  L.  Sp.  PI.  361.     1753. 
Coll.  :   Lyon  494,  Jefferson.     Aug. 

Polygonum  punctatum  ELL.  Bot.  S.  C.  &  Ga.  i :  455.    1817. 
Coll.  :  Wheeler  537,  539,  Jefferson.     Aug. 

Polygonum  orientale  L.  Sp.  PI.  362.     1753. 
Coll.  :  Wheeler  448,  Jefferson.     Aug. 


Wheeler :   FLORA  OF  SOUTHEASTERN  MINNESOTA.          385 

Polygonum  virginianum  L.  Sp.  PI.  360.     1753. 

Coll.  :  Wheeler  580,  Jefferson;  597. 
Crooked  Creek.     Aug. 
The  only  previous  collections  from  Minnesota  are  Sheldon, 

Madison  lake  and  Sheldon,  Zumbrota.     Infrequent  in  moist 

woods  along  Mississippi  river. 
Polygonum  ramosissimumMicHx.  Fl.  Bor.  Am.  i :  237.     1803. 

Coll.:  Wheeler  514,  Winnebago  ;  531,   Jefferson.     Aug. 
Polygonum  tenue  MICHX.  Fl.  Bor.  Am.  i :  238.     1803. 

Coll.  :  Wheeler,  351,  Crooked  Creek.     July. 
The  only   previous  authentic   collection    from   Minnesota  is 

Moyer,  Montevideo.     Infrequent  on  dry  sandy  ridges. 
Polygonum  convolvulus  L.  Sp.  PL  364.     1753. 

Coll.:   Wheeler  451,  Jefferson.     Aug. 
Polygonum  scandens  L.  Sp.  PL  364.     1753. 

Coll.  :  Wheeler  646,  Jefferson.     Aug. 
Polygonum  sagittatum  L.  Sp.  PL  363.     1753. 

Coll.  :  Wheeler  387,  Jefferson.     July. 

CHENOPODIACE^E. 

Chenopodium  botrys  L.  Sp.  PL  219.     1753. 

Coll.  :  Lyon  472,  Jefferson.     Aug. 
Salsola  tragus  L.  Sp.  PL  Ed.  2,  322.     1762. 

Coll.  :   Lyon  396,  Jefferson.     Aug. 

AMARANTHACE^E. 

Amaranthus  retroflexus  L.  Sp.  PL  991.     1753. 

Coll.  :  Wheeler  598,  Crooked  Creek.     Aug. 
Amaranthus  blitoides  S.  WATS.  Proc.  Am.  Acad.  12:   273. 
1877. 

Coll.  :  Wheeler  452,  Jefferson.     Aug. 
Acnida  tamariscina  (NUTT.)  WOOD,  Bot.  &  Fl.  289.     1873. 

Coll.  :  Wheeler  522,  547,  Jefferson.     Aug. 
Froelichia  floridana  (NUTT.)  Moo^  in  DC.  Prodr.  13:  Part  2, 
420.     1849. 

Coll.  :   Lyon  304,  325,  Jefferson.     Aug. 

NYCTAGINACE^. 

Allionia  nyctaginea  MICHX.  FL  Bor.  Am.  i :  100.     1803. 
Coll.  :  Wheeler  76,  Lyon  40,  Winnebago.     June. 


386  MINNESOTA    BOTANICAL    STUDIES. 

Allionia  linearis  PURSH,  Fl.  Am.  Sept.  728.     1814. 

Coll.  :   Wheeler  392,  Jefferson.     July. 

Not  previously  collected  in  Minnesota.  Collections  Oest- 
lund  155  and  Herrick  256,  Minneapolis,  in  the  Herbarium 
of  the  University  and  reported  in  Metaspermae  of  Minne- 
sota Valley  as  A.  linearis  Pursh  ?  should  be  A,  hirsuta 
Pursh. 

AIZOACE^E. 

Mollugo  verticillata  L.  Sp.  PL  89.     1753. 

Coll.  :    Lyon  279,  302,  Jefferson.     July,  Aug. 

CARYOPHYLLACE^E. 

Silene  stellata  (L.)  AIT.  f.  Hort.  Kew.  3:  84.     1811. 

Coll.  :  Lyon  254,  Wheeler  508,  Winnebago.     July,  Aug. 

Silene  alba  MUHL.  Cat.  45.     1813. 

Coll.  :  Lyon  164,  Winnebago.     June. 

The  only  previous  collections  from  Minnesota  are  from 
Goodhue  and  Winona  counties.  Frequent  along  Winne- 
bago and  Crooked  creeks. 

Silene  antirrhina  L.  Sp.  PL  419.     1753. 

Coll.  :  Wheeler  171,  Winnebago.     June. 

Cerastium  longipedtmculatum  MUHL.  Cat.  46.     1813. 
Coll.  :  Lyon  8,  Winnebago.     June. 

NYMPHjEACE^E. 

Nymphaea  advena  SOLAND.  in  Ait.  Hort.  Kew.  2  :  226.     1789. 
Coll.:    Wheeler    307,    Crooked  Creek;     454,   Jefferson. 

July,  Aug.     (Plate  XXVII.,  B.) 
Castalia  tuberosa    (PAINE)    GREENE,  Bull.  Torr.    Club,  15: 

84.     1888. 

Coll.  :  Wheeler  293,  439,  496,  Jefferson.     July,  Aug. 
Nelumbo  lutea  (WILLD.)  PERS.  Syn.  i :  92.     1805. 

Coll.  :  Wheeler  409,  494,  Jefferson.     July,  Aug. 
Abundant  in  the  sloughs  of  the  Mississippi  river  at  Jefferson. 
(Plate  XXV.,  A.) 

CERATOPHYLLACE^E. 

Ceratophyllum  demersum  L.  Sp.  PL  992.     1753. 

Coll.  :   Lyon  367,  480,  Jefferson.     Aug. 
Common  in  the  sloughs  of  the  Mississippi   river  at  Jefferson 
and  Crooked  Creek. 


Wheeler:   FLORA  OF  SOUTHEASTERN  MINNESOTA.  387 

RANUNCULACE.E. 

Caltha  palustris  L.  Sp.  PL  558.     1753. 

Coll.  :  Lyon  244,  Crooked  Creek.     July. 
Actsea  rubra  (AiT.)  WILLD.  Enum.  561.     1809. 

Coll.:   Lyon  450,  Jefferson.     Aug. 
Actaea  alba  (L.)  MILL.  Gard.  Diet.  Ed.  8,  No.  2.     1768. 

Coll.  :  Lyon  16,  Winnebago.     June. 
Aquilegia  canadensis  L.  Sp.  PL  533.     1753. 

Coll.  :  Wheeler  84,  Winnebago.     June. 

Anemone  virginiana  L.  Sp.  PL  540.     1753. 

Coll.  :  Lyon  9,  Winnebago;   245,  Crooked  Creek.     June, 

J^y. 

Anemone  canadensis  L.  Syst.  Ed.  12,  3:  App.  231.     1768. 
Coll.:  Lyon  286*^,  Jefferson.     July. 

Anemone  quinquefolia  L.  Sp.  PL  541.     1753. 

Coll.  :  Herb.  Wheeler  4,  Winnebago.     May. 

Hepatica  acuta  (PURSH)   BRITTON,  Ann.  N.  Y.  Acad.  Sci.  6: 

234.     1891. 
Coll.  :  Wheeler  134,  Winnebago.     June. 

Syndesmon  thalictroides  (L.)  HOFFMG.   Flora,   15:    Part  2, 

Intell.  Bl.  4,  34.      1832. 
Coll.  :   Wheeler  36,  Winnebago,     June. 

Pulsatilla  hirsutissima  (PURSH)  BRITTON,  Am.  N.  Y.  Acad. 

Sci.  6  :   217.     1891. 
Coll.  :  Wheeler  73,  Winnebago.     June. 

Clematis  virginiana  L.  Amoen.  Acad.  4:  275.     1759. 

Coll.:    Wheeler  194,  Winnebago;    355,  Crooked  Creek. 

June,  July. 
Atragene  americana  SIMS,  Bot.  Mag.  pL  887.     1806. 

Coll.:  Wheeler  259,  Winnebago ;  320,  Mayville.     June, 

July. 

Ranunculus  delphinifolius  TORR.  ;    Eaton,  Man.  Ed.   2,395. 

1818. 
Coll.  :  Lyon  201,  Crooked  Creek.     July. 

Ranunculus  abortivus  L.  Sp.  PL  551.     1753. 
Coll.  :  Wheeler  15,  Winnebago.     June. 

Ranunculus  pennsylvanicus  L.  f.  Suppl.  272.     1781. 
Coll.  :   Lyon  364,  378,  Jefferson.     Aug. 


388  MINNESOTA    BOTANICAL    STUDIES. 

Ranunculus  septentrionalis  Pom.   in  Lam.   Encycl.  6:    125. 
1804. 

Coll.  :  Wheeler  6,  Winnebago.     June. 

Batrachium  trichophyllum  (CHAIX)  BOSSCH,  Prodr.  Fl.  Bot. 
5-     1850. 

Coll.  :  Lyon  67,  Winnebago.     June. 
Batrachium  divaricatum  (  SCHRANK  )  WIMM.  Fl.  Schles.     1841 . 

Coll.  :  Lyon  219,  Mayville.     July. 
Thalictrum  dioicum  L.  Sp.  PI.  545.     1753. 

Coll.  :  Herb.  Wheeler  13,  Winnebago.     June. 
Thalictrum  purpurascens  L.  Sp.  PL  546.     1753. 

Coll.  :  Lyon  99,  Winnebago.     June. 

BERBERIDACE^E. 
Caulophyllum  thalictroides  (L.)  MICHX.  Fl.  Bor.  Am.  i :  205. 

1803. 

Coll.  :  Lyon  92,  Winnebago.     June. 
Podophyllum  peltatum  L.  Sp.  PI.  505.     1753. 

Coll.  :   Lyon  i,  Wheeler  157,  Winnebago.     June. 

MENISPERMACE^E. 

Menispermum  canadense  L.  Sp.  PL  340.     1753. 
Coll.  :   Wheeler  188,  Winnebago.     June. 

PAPAVERACE^:. 

Sanguinaria  canadensis  L.  Sp.  PL  505.     1753. 
Coll.  :    Lyon  169,  Crooked  Creek.     June. 
Bicuculla  cucullaria  (L.)  MILLSP.  Bull.  W.   Va.    Agric.  Exp. 

Sta.  2  :  327.      1892. 
Coll.  :   Wheeler  12,  Winnebago.     June. 

CRUCIFERJE. 
Lepidium  apetalum  WILLD.  Sp.  PL  3:   439.     1801. 

Coll.  :    Lyon  123,  Winnebago.     June. 
Sisymbrium  officinale  (L.)  SCOP.  FL  Cam.  Ed.  2,  2  :   26.     1772. 

Coll.  :   Lyon  422,  Winnebago.     Aug. 
Sisymbrium  altissimum  L.  Sp.  PL  659.     1753. 

Coll.  :  Lyon  273,  Wheeler  481,  Jefferson.     July,  Aug. 
Brassica  nigra  (L.)  KOCH,  in  Roehl,  Deutsche  Fl.  Ed.  3,  4: 
7i3-      1833. 

Coll.  :  Lyon  233,  Crooked   Creek.     July. 


\\7iecler:   FLORA  OF  SOUTHEASTERN  MINNESOTA.          389 

Brassica  arvensis  (L.)  B.S.P.  Prel.  Cat.  N.  Y.     1888. 

Coll.  :  Lyon  86,  Winnebago.     June. 
Roripa  palustris  (L.)  BESS.  Enum.  27.     1821. 

Coll.  :  Lyon  200,  Crooked  Creek.     July. 

Roripa  nasturtium  (L.)  RUSBY,  Mem.  Torr.  Club,  3:  Part  3, 
5-      1893. 

Coll.  :  Lyon  89,  Winnebago.     June. 

Cardamine   bulbosa   (SCHREB.)   B.S.P.    Prel.    Cat.   N.  Y.   4. 
1888. 

Coll. :  Wheeler  167,  Winnebago.     June. 
Bursa  bursa-pastoris  (L.)  BRITTON,  Mem.  Torr.  Club,  5  :  172. 
1894. 

Coll.  :  Lyon  120,  Winnebago.     June. 
Camelina  sativa  (L.)  CRANTZ,  Stirp.  Austr.  i :  18.     1762. 

Coll.  :  Lyon  213,  Mayville.     July. 
Draba  caroliniana  WALT.  Fl.  Car.  174.     1788. 

Coll.  :  Lyon  47,  Winnebago.     June. 

Arabis  laevigata  (MuHL.)  POIR.  in  Lam.  Encycl.  Suppl.  i :  411. 
1810. 

Coll.  :  Wheeler  154,  Winnebago.     June. 
Arabis  canadensis  L.  Sp.  PL  665.     1753. 

Coll.:     Lyon    in,    Winnebago;    212,    Crooked    Creek. 

June,  July. 
Arabis  glabra  (L.)  BERNH.  Verz.  Syst.  Erf.  195.     1800. 

Coll.:  Lyon  122,  Winnebago  ;  227,  Mayville.    June,  July. 
Erysimum  cheiranthoides  L.  Sp.  PI.  66 1.     1753. 

Coll.:   Lyon  187,  Crooked  Creek ;   226,  Mayville.     July. 

CAPPARIDACE^E. 

Polanisia  graveolens  RAF.  Am.  Journ.  Sci.  i :  378.     1819. 
Coll.  :   Lyon  277,  Jefferson.     July. 

CRASSULACE^E. 

Penthorum  sedoides  L.  Sp.  PI.  432.     1753. 
Coll.  :  Wheeler  384,  Jefferson.     July. 

SAXIFRAGACE^E. 

Saxifraga  pennsylvanica  L.  Sp.  PI.  399.     1753. 

Coll.  :  Wheeler  180,  Winnebago.     June. 
Heuchera  hispida  PURSH,  Fl.  Am.  Sept.  188.     1814. 

Coll.  :  Wheeler  83,  Winnebago.     June. 


390  MINNESOTA    BOTANICAL    STUDIES. 

Mitella  diphylla  L.  Sp.  PL  406.     1753. 

Coll.  :  Wheeler  40,  Winnebago.     June. 
Parnassia  caroliniana  MICHX.  Fl.  Bor.  Am.  i :   184.     1803. 
Coll.:   Wheeler  587,    Crooked  Creek;   629,  Brownsville. 
Aug. 

GROSSULARIACE^:. 

Ribes  cynosbati  L.  Sp.  PI.  202.     1753. 

Coll.  :   Wheeler  77,  no,  Winnebago.     June. 
Ribes  uva-crispa  L.  Sp.  PI.  201.     1753. 

Coll.:   Wheeler  30,  75,  248,  Winnebago.     June. 
Frequently  adventive  in  open  woods  throughout. 
Ribes  floridum  L'HER.  Stirp.  Nov.  i :   4.     1784. 

Coll.  :   Lyon  82,  Winnebago.    June. 

HAMAMELIDACE^:. 

Hamamelis  virginiana  L.  Sp.  PI.  124.     1753. 
Coll.  :  Wheeler  150,  Winnebago.     June. 
Reported  from  southeastern  Winona  County.     No  Minnesota 
specimens  in  the  Herbarium  of  the  University.     Local  on 
north  side  of  bluff  in  section  22  of  Winnebago. 

ROSACE^E. 

Opulaster  opulifolius~(L.)  KUNTZE,  Rev.  Gen.  PL  949.     1891. 

Coll.  :  Lyon  33,  103,  Winnebago.  June. 
Spiraea  salicifolia  L.  Sp.  PL  489.  1753. 

Coll.  :  Lyon  438,  464,  Jefferson.     Aug. 
Rubus  occidentalis  L.  Sp.  PL  493.     1753. 

Coll.:    Lyon    55,   Winnebago;    Wheeler  453,  Jefferson. 

June,  Aug. 
Rubus  villosus  AIT.  Hort.  Kew.  2:  210.     1789. 

Coll.  :  Wheeler  105,  Lyon  296,  Winnebago.     June,  Aug. 
Rubus  canadensis  L.  Sp.  PL  494.     1753. 

Coll.:    Wheeler  396,  Jefferson.     July. 
Fragaria  americana  (PORTER)  BRITTON,  Bull.  Torr.  Club,  19 : 

222.       1892. 

Coll.  :  Wheeler  135,  Winnebago.  June. 
Potentilla  arguta  PURSH,  Fl.  Am.  Sept.  736.  1814. 

Coll.  :  Lyon  247,  Crooked  Creek.  July. 
Potentilla  monspeliensis  L.  Sp.  PL  499.  1753. 

Coll.  :  Lyon  439,  Jefferson.     Aug. 


Wheeler:   FLORA  OF  SOUTHEASTERN  MINNESOTA.  391 

Potentilla  canadensis  L.  Sp.  PL  498.     1753. 
Coll.  :  Wheeler  86,  Winnebago.     June. 

Geum  canadense  JACC^  Hort.  Vind.  2:  82.  pi,  775.     1772. 
Coll.  :   Lyon  209,  Crooked  Creek.     July. 

Geum  strictum  AIT.  Hort.  Kew.  2:  217.     1789. 

Coll.  :  Wheeler   277,   Lyon   195,   Crooked  Creek.     June, 


Agrimonia  hirsuta  (MUHL.)  BICKNELL,  Bull.  Torr.  Club,  23  : 

509.     1896. 
Coll.  :  Lyon  196,  228,  Crooked  Creek.     July. 

Rose  blanda  AIT.  Hort.  Kew.  2:  202.     1789. 

Coll.  :  Lyon  37,  431,  Winnebago.     June,  Aug. 

Rosa  arkansana  PORTER,  Syn.  Fl.  Colo.  38.     1874. 

Coll.:    Wheeler  441,    Lyon    343,   Jefferson;  Lyon    339, 
Winnebago.     Aug. 

POMACEJE. 
Malus  ioensis  (Wooo)  BRITTON,  in  Britt.  &  Brown,  111.  Fl.  2  : 

235.7.  i98°-     l897- 

Coll.  :  Wheeler  88,  160,  Winnebago  ;  605,  Crooked  Creek. 
June,  Aug. 

Amelanchier  canadensis  (L.)  MEDIC.  Geschichte,  79.     1793. 
Coll.  :  Wheeler  37,  500,  Winnebago.     June,  Aug. 

Amelanchier  botryapium  (L.  f.)  DC.  Prodr.  2:  632.     1825. 
Coll.  :  Wheeler  253,  Winnebago.    June. 

Amelanchier  alnifolia  NUTT.  ;    T.  &  G.   Fl.  N.  A.  i  :  473. 

As  synonym.      1840. 
Coll.  :  Wheeler  203,  Winnebago.     June. 

Crataegus  punctata  JACO^  Hort.  Vind.  i:  10.  pi.  28.     1770. 
Coll.  :  Wheeler  141,  651,  Winnebago.    June,  Aug. 

Crataegus  coccinea  L.  Sp.  PL  476.     1753. 
Coll.  :  Lyon  101,  Winnebago,  June. 

Crataegus  macracantha  LODD.  ;  Loudon,  Arb.  Brit.  Ed.  2,  2: 

819.      1854. 

Coll.  :  Wheeler  499,  Winnebago.     Aug. 
Not  previously  reported  from  Minnesota. 

Crataegus  tomentosa  L.  Sp.  PL  476.     1753. 

Coll.  :  Lyon  3,  Wheeler  140,  Winnebago.     June. 


392  MINNESOTA    BOTANICAL    STUDIES. 

DRUPACE^E. 

Prunus  americana  MARSH.  Arb.  Am.  in.     1785. 
Coll.  :  Wheeler  353,  Crooked  Creek.    July. 

Prunus  nigra  AIT.  Hort.  Kew.  2  :   165.     1789. 

Coll.:    Wheeler   321,    354,    Mayville ;    501,    Winnebago. 

July,  Aug. 

Not  previously  reported  from  Minnesota.  This  species  has 
been  recognized  by  horticulturists  in  several  parts  of  the 
state  but  no  previous  authentic  collections  are  known  to 
have  been  made.  It  is  common  on  the  lowlands  of  the 
North  and  South  forks  of  Crooked  creek.  Also  col- 
lected on  the  banks  of  Winnebago  creek  and  in  East 
Burns  valley,  Winona  county. 

Prunus  virginiana  L.  Sp.  PI.  473.     1753. 

Coll.  :   Wheeler  345,  Crooked  Creek.     July. 

Prunus  serotina  EHRH.  Beitr.  3:  20.     1788. 
Coll.  :   Wheeler  178,  Winnebago.     June. 

C^ESALPINACEjE. 

Cassia  chamaecrista  L.  Sp.  PL  379.     1753. 
Coll.  :  Lyon  256,  Jefferson.     July. 

Gleditsia  triacanthos  L.  Sp.  PL  1056.     1753. 

Coll.  :  Lyon  and  Wheeler;  413,  W.  Jefferson.  July. 
Not  previously  reported  from  Minnesota.  This  tree  has  been 
frequently  cultivated  for  ornament  throughout  the  southern 
part  of  the  state  but  no  native  trees  have  previously  been 
reported.  It  is  frequent  on  the  islands  of  the  Mississippi 
river  in  northeastern  Iowa  and  extends  north  along  the 
river  into  Houston  county,  Minnesota,  where  it  probably 
reaches  its  northern  limit.  The  tree  from  which  the  col- 
lection was  made  is  59  feet  high  and  has  a  trunk-circum- 
ference of  6  feet,  3  feet  from  the  ground. 

Gymnocladus  dioica  KOCH,  Dendrol.  1:5.     1869. 

Coll.:    Lyon   193,   230,  Crooked   Creek ;    271,  Jefferson. 
July.     (Plate  XXIV.,  A.) 

PAPILIONACE^E. 

Baptisia  bracteata  ELL.  Bot.  S.  C.  &  Ga.  i :  469.     1817. 

Coll.:   Wheeler  94,   Lyon   34,   Winnebago;     Lyon    202, 
Crooked  Creek.     June,  July. 


Wheeler :    FLORA  OF  SOUTHEASTERN  MINNESOTA.  393 

Baptisia  leucantha  T.  &  G.  Fl.  N.  A.  i :  385.     1840. 

Coll.  :   Lyon  194,  Crooked  Creek.     July. 
Trifolium  hybridum  L.  Sp.  PI.  766.     1753. 

Coll.  :  Lyon  421,  Winnebago.     Aug. 
Trifolium  repens  L.  Sp.  PI.  767.     1753. 

Coll.  :  Lyon  118,  Winnebago.     June. 
Amorpha  fruticosa  L.  Sp.  PL  713.     1753. 

Coll.  :  Lyon  473,  Jefferson.     Aug. 
Amorpha  canescens  PURSH,  Fl.  Am.  Sept.  467.     1814. 

Coll.  :  Wheeler  291,  Mayville.     July. 

Kuhnistera   Candida  (WILLD.)   KUNTZE,    Rev.   Gen.  PI.  192. 

1891. 
Coll.  :  Wheeler  397,  Jefferson.     July. 

Kuhnistera  purpurea  (VENT.)  MAcM.  Met.  Minn.  329.     1892. 
Coll.  :   L}'on  262,  Jefferson.     July. 

Astragalus  carolinianus  L.  Sp.  PI.  757.     1753. 
Coll.  :    Lyon  257,  395,  Jefferson.     July,  Aug. 

Meibomia  grandiflora    (WALT.)  KUNTZE,  Rev.   Gen.  PI.   196. 

1891. 
Coll.  :  Lyon  198,  246,  Crooked  Creek.    July. 

Meibomia  dillenii  (DARL.)  KUNTZE,  Rev.  Gen.  PI.  195.    1891. 

Coll.  :   Wheeler  482,  Jefferson.     Aug. 

No  previously  collected   Minnesota  specimens   in  the  Her- 
barium of  the  University. 

Meibomia  illinoensis  (A.  GRAY)  KUNTZE,  Rev.   Gen.  PL   198. 

1891. 
Coll.  :    Wheeler    390,   446,  Jefferson ;    609,    Brownsville. 

July,  Aug. 

Not  previously  reported  from  Minnesota.     Frequent  and   in 
some  places  common  in  dry  fields  and  hillsides. 

Meibomia  canadensis  (L.)  KUNTZE,  Rev.  Gen.  PL  195.     1891. 
Coll.  :   Wheeler  331,  Crooked  Creek.     July. 

Lespedeza  capitata  MICHX.  Fl.  Bor.  Am.  2:  71.     1803. 
Coll.  :  Wheeler  525,  Jefferson.     Aug. 

Lathyrus  venosus  MUHL.  ;  Willd.  Sp.  PL  3:  1092.     1803. 
Coll.  :  Lyon  48,  Winnebago.     June. 

Lathyrus  ochroleucus  HOOK.  FL  Bor.  Am.  i :   159.     1833. 
Coll.  :  Lyon  87,  Winnebago.     June. 


394  MINNESOTA    BOTANICAL    STUDIES. 

Falcata  comosa  (L.)  KUNTZE,  Rev.  Gen.  PL  182.     1891. 

Coll.:  Wheeler  507,   Lyon   332,  Winnebago ;  Lyon  469, 

Jefferson.     Aug. 
Falcata  pitcher!  (T.  &  G.)  KUNTZE,  Rev.  Gen.  PL  182.     1891. 

Coll.  :  Wheeler  573,  Jefferson.     Aug. 

Not  previously  reported  from  Minnesota.     Several  specimens 
of  this   species,    previously  reported  as  f.   comosa,  have 
been  collected  in  southern  Minnesota.     Probably  common 
throughout  the  southern  part  of  the  state. 
Apios  apios  (L.)  MACM.  Bull.  Torr.  Club,  19:   15.     1892. 

Coll.  :  Wheeler,  337,  Crooked  Creek  ;  Wheeler  399,  Lyon 

388,  Jefferson.     July,  Aug. 
Strophostyles  helvola  (L.)  BRITTOX  in  Britt.  &  Brown,  111.  Fl. 

2:  338.7.  2235.     1897. 
Coll.  :  Lyon  387,  Jefferson.     Aug. 

GERANIACE^E. 

Geranium  maculatum  L.  Sp.  PL  68 1.     1753. 
Coll.  :  Lyon  4,  Winnebago.     June. 

OXALIDACE^E. 

Oxalis  violacea  L.  Sp.  PL  434.     1753. 

Coll.  :  Lyon  50,  Winnebago.     June. 
Oxalis  stricta  L.  Sp.  PL  435.     1753. 

Coll.  :  Lyon  81,  Winnebago.     June. 

LINAGES. 

Linum  sulcatum  RIDDEL,  Suppl.  Cat.  Ohio  PL  10.     1836. 
Coll.  :  Wheeler  290,  Mayville  ;  431,  Jefferson.     July. 

RUTACEJE. 

Xanthoxylum  americanum  MILL.   Gard.   Diet.   Ed.  8,  No.  2. 

1768. 
Coll.  :  Wheeler  132,  413,  Winnebago.     June,  Aug. 

POLYGALACE^E. 

Polygala  verticillata  L.  Sp.  PL  706.     1753. 

Coll.:     Wheeler   342,    Crooked    Creek;    428,   Jefferson. 

July. 

Polygala  viridescens  L.  Sp.  PL  705.     1753. 
Coll.  :   Lyon  483,  Jefferson.     Aug. 


Wheeler :   FLORA  OF  SOUTHEASTERN  MINNESOTA.          395 

Polygala  senega  L.  Sp.  PI.  704.     1753. 
Coll.  :  Lyon  21,  Winnebago.     June. 

EUPHORBIACE^E. 

Acalypha  virginica  L.  Sp.  PI.  1003.     1753. 

Coll.  :  Lyon  349,  397,  463,  Jefferson.     Aug. 
Euphorbia  glyptosperma  ENGELM.   Bot.   Mex.  Bound.  Surv. 
187.     1859. 

Coll.  :  Wheeler  434,  Jefferson.     July. 
Euphorbia  maculata  L.  Sp.  PI.  455.     1753. 

Coll.  :  Wheeler  430,  Jefferson.     July. 
Euphorbia  nutans  LAG.  Gen.  &  St.  17.     1816. 

Coll.  :  Wheeler  336,  Crooked  Creek.     July. 
Euphorbia  corollata  L.  Sp.  PI.  459.     1753. 

Coll.  :   Lyon  189,  Wheeler  375,  Crooked  Creek.     July. 
Euphorbia  heterophylla  L.  Sp.  PI.  45 3.     1753. 

Coll.  :  Wheeler  466,  Jefferson.     Aug. 
Euphorbia  cyparissias  L.  Sp.  PI.  461.     1753. 

Coll.  :   Lyon  437,  Jefferson.     Aug. 

ANACARDIACEJE. 

Rhus  hirta  (L.)  SUDW.  Bull.  Torr.  Club,  19  :  82.     1892. 

Coll.:  Lyon  319, Jefferson.  Aug. 
Rhus  glabra  L.  Sp.  PI.  265.  1753. 

Coll.  :  Lyon  272,  Jefferson.  July. 
Rhus  radicans  L.  Sp.  PI.  266.  1753. 

Coll.  :  Lyon  327,  350,  Jefferson.     Aug. 

CELASTRACEJE. 

Euonymus   atropurpureus  JACQ^  Hort.  Vind.  2:    5.  pi.  120. 

1772. 
Coll.:    Lyon    140,    Winnebago;     263,   Jefferson.     June, 

July. 
Celastrus  scandens  L.  Sp.  PI.  196.     1753. 

Coll.:   Wheeler   104,  Winnebago;    Lyon  380,  Jefferson. 
June,  Aug. 

STAPHYLEACE^E. 

Staphylea  trifolia  L.  Sp.  PL  270.     1753. 

Coll.  :  Wheeler  202,  Winnebago.     June. 


396  MINNESOTA    BOTANICAL    STUDIES. 

ACERACE^:. 

Acer  saccharinum  L.  Sp.  PI.  1055.     1753. 

Coll.:     Lyon    149,   Winnebago  ;     274,   Jefferson.     June, 


Acer  nigrum  MICHX.  f.  Hist.  Arb.  Am.  2  :   238.  pL  16.     1810. 
Coll.  :  Wheeler  149,  Winnebago.     June. 

Acer  spicatum  LAM.  Encycl.  2:  381.     1786. 

Coll.:    Wheeler    163,   198,  Winnebago;    319,    May  ville  ; 

625,  Brownsville.     June,  Aug. 
Frequent  on  moist  shaded  cliffs  throughout. 

Acer  negundo  L.  Sp.  PI.  1056.     1753. 

Coll.:  Lyon  119,  Winnebago.     June. 

BALSAMINACE^E. 

Impatiens  aurea  MUHL.  Cat.  26.     1813. 
Coll.  :  Wheeler  328,  Mayville.     July. 

RHAMNACEJE. 

Ceanothus  americanus  L.  Sp.  PL  195.     1753. 

Coll.  :    Lyon    66,    Winnebago  ;    Wheeler    356,    Crooked 
Creek.     June,  July. 

Ceanothus  ovatus  DESF.  Hist.  Arb.  2  :  381.     1809. 

Coll.  :   Wheeler  92,  Lyon  66^,  Winnebago.     June. 

VITACEiE. 

Vitis  vulpina  L.  Sp.  PL  203.     1753. 

Coll.:   Wheeler  139,  Winnebago;    344,  Crooked  Creek. 
June,  July. 

Parthenocissus  quinquefolia  (L.)  PLANCH,  in  DC.  Mon.  Phan. 

5:  Part  2,  448.     1887. 

Coll.:  Wheeler  235,  Winnebago;    Lyon  379,    Jefferson. 
June,  Aug. 

TILIACE^E. 

Tilia  americana  L.  Sp.  PL  514.     1753. 
Coll.  :  Lyon  211,  Mayville.     July. 

MALVACEAE. 

Malva  rotundifolia  L.  S  p.  PL  688.     1753. 
Coll.:   Lyon  491,  Jefferson.     Aug. 


Wheeler:    FLORA  OF  SOUTHEASTERN  MINNESOTA.  397 

Napaea  dioica  L.  Sp.  PL  686.     1753. 

Coll.  :  Lyon  266,  Jefferson.     July. 

Previously  collected  at  Vasa  and  Lanesboro.     Rare  on  low- 
land near  Winnebago  creek. 

Abutilon   abutilon   (L.)    RUSBY,   Mem.  Torr.   Club,   5:    222. 

1894. 
Coll.  :  Lyon  283,  Jefferson.     July. 

HYPERICACE^:. 

Hypericum  ascyron  L.  Sp.  PI.  783.     1753. 

Coll.:     Lyon    235,    Crooked    Creek;    436,    Winnebago. 
June,  Aug. 

Hypericum  maculatum  WALT.  Fl.  Car.  189.     1788. 

Coll.:    Wheeler  442,  483,  Jefferson;    615,  Brownsville; 
Lyon  452,  Jefferson.     Aug. 

Hypericum  majus  (A.  GRAY)  BRITTON,  Mem.  Torr.  Club,  5 : 

225.     1894. 

Coll.:    Wheeler   427,   Jefferson;    601,    Crooked    Creek. 
July,  Aug. 

CISTACE^:. 

Helianthemum  majus  (L.)  B.S.P.  Prel.  Cat.  N.  Y.  6.     1888. 
Coll.:  Lyon   167,  Crooked   Creek;  Wheeler  477,  Jeffer- 
son;  633,  Brownsville.     Aug. 

Helianthemum  canadense  (L.)  MICHX.  Fl.  Bor.  Am.  i :  308. 

1803. 
Coll.  :  Wheeler  631,  Brownsville.     Aug. 

Lechea  stricta   LEGGETT;   BRITTON,   Bull.  Torr.  Club,   21: 

251.     1894. 

Coll.  :     Wheeler  359,  Crooked  Creek;  Lyon  331,  Winne- 
bago.    July,  Aug. 

VIOLACEJE. 

Viola  pedatifida  DON,  Gard.  Diet,  i :  320.     1831. 
Coll.  :  Wheeler  429,  Jefferson.     July. 

Viola  obliqua  HILL,  Hort.  Kew.  316.  pi.  12.     1769. 
Coll.:  Wheeler  114,  Winnebago.     June. 

Viola  pedata  L.  Sp.  PI.  933.     1753. 

Coll.  :  Wheeler  216,  Jefferson.     June. 


398  MINNESOTA    BOTANICAL    STUDIES. 

Viola  pubescens  AIT.  Hort.  Kevv.  3:  290.     1789. 
Coll.  :  Wheeler  205,  Winnebago.     June. 

THYMELEACE^E. 

Dirca  palustris  L.  Sp.  PL  358.     1753. 

Coll.  :  Wheeler  520,  Winnebago.     Aug. 
Rare  in  moist  thickets  near  Winnebago  creek. 

LYTHRACE^E. 

Lythrum  alatum  PURSH,  Fl.  Am.  Sept.  334.     1814. 
Coll.  :  Lyon  281,  Wheeler  416,  Jefferson.     July. 

ONAGRACEjE. 
Epilobium  coloratum  MUHL.  ;  Willd.  Enum.  i :  411.     1809. 

Coll.  :  Wheeler    479,  Jefferson  ;   608,  Brownsville.     Aug. 
Epilobium   adenocaulon   HAUSSK.    Oest.  Bot.    Zeit.    29:   119. 

1879. 
Coll.  :  Wheeler  323,  Mayville  ;  595,  Crooked  Creek  ;  Lyon 

457,  Jefferson.     July,  Aug. 

Onagra  biennis  (L.)  SCOP.  Fl.  Cam.  Ed.  2,  i:  269.     1772. 
Coll.:    Lyon  166^,  Winnebago;    286,  Jefferson.     June, 

July. 

(Enothera  rhombipetala  NUTT.  ;  T.  &  G.  Fl.  N.  A.  i :  493. 

1840. 
Coll.  :  Lyon  323,  Jefferson.     Aug. 

Gaura  biennis  L.  Sp.  PL  347.     1753. 

Coll.  :  Wheeler  574,  Jefferson.     Aug. 

No  previous  authentic  collection  from  Minnesota.  There  are 
no  Minnesota  specimens  in  the  Herbarium  of  the  Univer- 
sity. Miss  Manning's  collection  of  1883  from  Pepin,  Wis., 
is  probably  the  one  upon  which  is  based  the  report  of  this 
species  by  Upham  and  others. 

Circaea  lutetiana  L.  Sp.  PL  9.     1753. 

Coll.  :  Wheeler  270,  Crooked  Creek.     June. 

Circaea  alpina  L.  Sp.  PL  9.     1753. 

Coll.  :  Lyon  152,  Winnebago.     June. 

ARALIACE^E. 

Aralia  racemosa  L.  Sp.  PL  273.     1753. 
Coll.:  Lyon  345,  Jefferson.     Aug. 


\\'hcler :   FLORA  OF  SOUTHEASTERN  MINNESOTA.          399 

Aralia  nudicaulis  L.  Sp.  PI.  274.     1753. 
Coll.  :  Lyon  15,  Winnebago.     June. 
Panax  quinquefolium  L.  Sp.  PI.  1058.     1753. 

Coll.  :   Lyon  210,  Mayville  ;  Wheeler  469,  Jefferson.    July, 
Aug. 

UMBELLIFER^E. 

Angelica  atropurpurea  L.  Sp.  PI.  251.     1753. 

Coll.  :  Wheeler  311,  Crooked  Creek.     July. 
Heracleum  lanatum  MICHX.  Fl.  Bor.  Am.  i :   166.     1803. 

Coll.  :  Lyon  93,  Winnebago.     June. 
Sanicula  marylandica  L.  Sp.  PI.  235.     1753. 

Coll.:  Wheeler  175,  Winnebago.     June. 

Sanicula    gregaria    BICKNELL,    Bull.    Torr.    Club,    22 :    354. 

1895. 

Coll.  :  Wheeler  177,  Winnebago.     June. 
The  only  precious  collection  from  Minnesota  is  that  of   Shel- 
don, Milaca,   1892. 

Sanicula  canadensis  L.  Sp.  PL  235.     1753. 
Coll.  :  Lyon  260,  Jefferson.     July. 

Sanicula   trifoliata    BICKNELL,    Bull.  Torr.    Club,    22 :    359. 

1895. 

Coll.  :  Lyon  214,  Mayville.     July. 
Not  previously  reported  from  Minnesota. 

Pimpinella  integerrima  (L.)  A.  GRAY,  Proc.  Am.  Acad.  7 : 

345-     1868. 
.  Coll.  :  Wheeler  179,  Winnebago.     June. 

Washingtonia  claytoni  (Micnx.)  BRITTON  in  Brit.  &  Brown, 

111.  Fl.  2:  530./.  2680.     1897. 
Coll.  :  Lyon  7,  Winnebago.     June. 

Sium  cicutaefolium  GMEL.  Syst.  2:  482.     1791. 

Coll.  :  Wheeler  545,  Lyon  449,  Jefferson.     Aug. 

Zizia  aurea  (L.)  KOCH,  Nov.  Act.  Caes.  Leop.  12  :   129.     1825. 
Coll.:  Wheeler   174,   Winnebago;  Lyon   261,  Jefferson. 
June,  July. 

Zizia  cordata  DC.  Prodr.  4:  100.     1830. 
Coll.  :  Lyon  292,  352,  Jefferson.     July. 

Cicuta  maculata  L.  Sp.  PI.  256.     1753. 

Coll.  :  Wheeler  338,  Crooked  Creek.     July. 


400  MINNESOTA    BOTANICAL    STUDIES. 

Cicuta  bulbifera  L.  Sp.  PL  255.     1753. 

Coll.  :  Wheeler  607,  Brownsville.     Aug. 

Deringa  canadensis  (L.)  KUNTZE,  Rev.  Gen.  PI.  266.     1891. 
Coll.  :  Wheeler  193,  Winnebago.     June. 

Berula  erecta  (Huos.)   COVILLE,  Contr.   Nat.   Herb.  4 :    115. 

1893. 

Coll.  :  Wheeler  588,  Crooked  Creek.     Aug. 
In  cold  springs  at  the  head  of  Clear  creek. 

Hydrocotyle  americana  L.  Sp.  PI.  234.     1753. 

Coll.  :  Wheeler  314,  Mayville.     July. 

The  only  previously  reported  locality  of  collection  is   St. 
Croix  Falls.     Rare  in  moist  woods  near  Crooked   creek. 

CORNACE^E. 

Cornus  circinata  L'HER.  Cornus,  7.  pi.  j.     1788. 

Coll.  :  Wheeler  81,  Winnebago.     June. 
Cornus  amonum  MILL.  Gard.  Diet.  Ed.  8,  No.  5.     1768. 

Coll.  :  Lyon  351,  Jefferson.     Aug. 
Cornus  stolonifera  MICHX.  Fl.  Bor.  Am.  i :  92.     1803. 

Coll.  :  Wheeler  69,  173,  Winnebago.     June. 
Cornus  candidissima  MARSH,  Arb.  Am.  35.     1785. 

Coll.  :  Wheeler  172,  Winnebago.     June. 

PYROLACE^E. 

Pyrola  elliptica  NUTT.  Gen.  i :  273.     1818. 

Coll.:  Wheeler   191,   Winnebago;    276,  Crooked  Creek. 
June,  July. 

ERICACEAE. 

Arctostaphylos  uva-ursi  (L.)  SPRENG.  Syst.  2:  287.     1825. 

Coll.  :  Lyon  116,  Jefferson.     June. 
On  a  sandy  point  of  a  bluff  in  section  19  of  Jefferson. 

PRIMULACE^E. 

Lysimachia  terrestris  (L.)  B.S.P.  Prel.  Cat.  N.  Y.  34.     1888. 

Coll.  :  Lyon  249,  Crooked  Creek.  July. 
Steironema  ciliatum  (L.)  RAF.  Ann.  Gen.  Phys.  7  :  192.  1820. 

Coll.  :  Lyon  251,  Crooked  Creek.  July. 
Dodecatheon  meadia  L.  Sp.  PI.  144.  1753. 

Coll.  :  Wheeler  340,  Crooked  Creek.     July. 


Wheeler:   FLORA  OF  SOUTHEASTERN  MINNESOTA.          401 

Previously  collected   only  in  Winona  and  Wabasha  counties. 
Rare  in  moist  woods. 

OLEACE.E. 

Fraxinus  lanceolata  BORCK.  Handb.  Forst.  Bot.  i :  826.     1800. 

Coll.:  Lyon  300, Jefferson.     Aug. 
Fraxinus  nigra  MARSH.  Arb.  Am.  51.     1785. 

Coll.  :  Lyon  173,  Crooked  Creek.     June.    (Plate  XXIII., 
B.) 

GENTIANACE.E. 

Gentiana  crinita  FROEL.  Gen.  112.     1796. 

Coll.  :  Lyon  454,  488,  Jefferson.     Aug. 
Gentiana  quinquefolia  L.  Sp.  PI.  230.     1753. 
Coll.  :  Lyon  455,  487,  Jefferson.     Aug. 
Gentiana  flavida  A.  GRAY,  Journ.  Sci.  (II.)  i:  180.     1846. 
Coll.:  Wheeler  516,   Winnebago ;  596,  Crooked  Creek. 
Aug. 

APOCYNACE.E. 

Apocynum  androsaemifolium  L.  Sp.  PL  213.     1753. 

Coll.  :  Lyon  188,  Crooked  Creek.     July. 
Apocynum  cannabinum  L.  Sp.  PI.  213.     1753. 

Coll.  :  Lyon  471,  Jefferson.     Aug. 
Apocynum  cannabinum  glaberrimum  DC.  Prodr.  8  :  439.     1844. 

Coll.  :  Lyon  357,  Jefferson.     Aug. 

ASCLEPIADACE^. 

Asclepias  tuberosa  L.  Sp.  PI.  217.     1753. 

Coll.:  Wheeler   287,  369,   Crooked    Creek;  Lyon   356, 

Jefferson.     July,  Aug. 
Asclepias  incarnata  L.  Sp.  PI.  215.     1753. 

Coll.  :  Lyon  177,  Winnebago  ;  365,  Jefferson.    June,  Aug. 
Asclepias  obtusifolia  MICHX.  Fl.  Bor.  Am.  i:   115.     1803. 

Coll.  :  Wheeler  569,  Jefferson.     Aug. 
Previously  reported  only  by   Lapham.     Infrequent  on  dry 

hillsides. 
Asclepias  exaltata  (L.)  MUHL.  Cat.  28.     1813. 

Coll.  :  Lyon  178,  Winnebago.     July. 
Asclepias  syriaca  L.  Sp.  PI.  214.     1753- 

Coll.  :  Lyon  176,  Winnebago.     July. 


402  MINNESOTA    BOTANICAL    STUDIES. 

Asclepias  verticillata  L.  Sp.  PL  217.     1753. 

Coll.  :  Wheeler  286,  Crooked  Creek  ;  378,  Jefferson.    July. 
Acerates  viridiflora  (RAF.)  EATON,  Man.  Ed.  5,  90.     1829. 

Coll.:  Lyon    179,   Winnebago ;    309^,  Jefferson.     July, 
Aug. 

CONVOLVULACE.E. 

Convolvulus  sepium  L.  Sp.  PI.  153.     1753. 

Coll.  :  Wheeler  306,  Crooked  Creek.     July. 
Convolvulus  spithamaeus  L.  Sp.  PI.  158.     1753. 

Coll.:    Wheeler    207,    Winnebago;    358,    371,    Crooked 
Creek;  385,  Jefferson.     June,  July. 

CUSCUTACEjE. 

Cuscuta  indecora  CHOISY,  Mem.  Soc.  Gen.  9  :   278.  pi.  j.  f.  5. 

1841. 

Coll.  :  Wheeler  436,  557,  647,  Jefferson.     July,  Aug. 
Cuscuta   coryli  ENGELM.  Am.  Journ.  Sci.  43:    337.  f.  7-11. 

1842. 

Coll.  :  Wheeler  503,  Winnebago.     Aug. 
Cuscuta  gronovii  WILLD.  ;  R.  &  S.  Syst.  6  :   205.     1820. 

Coll.:    Wheeler    308,    592,    Crooked    Creek;    438,    440, 

Jefferson.     July,  Aug. 
Cuscuta  paradoxa  RAF.  Ann.  Nat.  13.     1820. 

Coll.  :  Wheeler  437,  648,  Jefferson.     July,  Aug. 

POLEMONIACE^E. 

Phlox  pilosa  L.  Sp.  PL  152.     1753. 

Coll.  :  Herb.  Wheeler  14,  Winnebago.     June. 
Phlox  divaricata  L.  Sp.  PL  152.     1753. 

Coll.  :  Lyon  5,  Winnebago.     June. 
Polemonium  reptans  L.  Syst.  Ed.  10,  No.  i.     1759. 

Coll.  :  Wheeler  33,  Winnebago.     June. 

HYDROPHYLLACE^E. 

Hydrophyllum  virginicum  L.  Sp.  PL  146.     1753. 

Coll.  :  Lyon  n,  Winnebago.     June. 
Hydrophyllum  appendiculatum  MICHX.  Fl.  Bor.  Am.  i :  134. 

1803. 
Coll.  :  Wheeler  324,  Mayville.     July. 


Wheeler :   FLORA  OF  SOUTHEASTERN  MINNESOTA.          403 

Macrocalyx  nyctelea  (L.)  KUNTZE,  Rev.  Gen.  PL  434.     1891. 
Coll.  :   Lyon  19,  Winnebago.     June. 

BORAGINACE^:. 

Lappula  lappula  (L.)  KARST.  Deutsch.  Fl.  979.      1880-83. 

Coll.  :  Lyon  186,  Crooked  Creek.     July. 
Lappula  virginianum  (  L.)  GREENE,  Pittonia,  2  :   182.     1891. 

Coll.  :  Lyon  237,  Mayville.     July. 

Lithospermum  gmelini  (  MICHX.)  A.  S.  HITCHCOCK,  Spring  Fl. 
Manh.  30.      1894. 

Coll.:  Lyon  281^,  Jefferson.     July. 
Lithospermum  canescens  (MICHX.)  LEHM.  Asperif.  305.     1818. 

Coll.  :   Lyon  27,  Winnebago.     June. 

Lithospermum  angustifolium   MICHX.   Fl.  Bor.   Am.    i :    130. 
1803. 

Coll.  :  Wheeler  450,  Jefferson.     Aug. 
Onosmodium  caroliniana  (LAM.)  DC.  Prodr.  10 :  70.     1846. 

Coll.  :  Wheeler  352,  Crooked  Creek.     July. 
Lycopsis  arvensis  L.  Sp.  PL  139.     1753. 

Coll.  :  Lyon  no,  Winnebago.     June. 
Not  previously  reported  from  Minnesota. 

VERBENACE^E. 

Verbena  urticifolia  L.  Sp.  PL  20.     1753. 

Coll.  :  Wheeler  406,  548,  Jefferson.     July,  Aug. 
Verbena  hastata  L.  Sp.  PL  20.     1753. 

Coll.  :  Wheeler  403,  Jefferson.     July. 
Verbena  stricta  VENT.  Desc.  PL  Jard.  Cels.  -pi.  53.     1800. 

Coll.  :  Wheeler  401,  Jeffepson.    July. 
Verbena  bracteosa  MICHX.  FL  Bor.  Am.  2  :  13.     1803. 

Coll.  :  Wheeler  635,  Brownsville.     Aug. 
Lippia  lanceolata  MICHX.  FL  Bor.  Am.  2  :  15.     1803. 

Coll.:    Lyon   279,  Jefferson;   Wheeler  622,  Brownsville. 

July,  Aug. 
Common  on  the  very  low  lands  of  the  Mississippi  river. 

LABIATE. 

Teucrium  canadense  L.  Sp.  PL  564.     1753. 
Coll.  :   Wheeler  414,  Jefferson.     July. 


404  MINNESOTA    BOTANICAL    STUDIES. 

Scutellaria  lateriflora  L.  Sp.  PI.  598-     I7S3- 

Coll.  :   Wheeler  455,  Jefferson  ;  517,  Winnebago.     Aug. 

Scutellaria  cordifolia  MUHL.  Cat.  56.     1813. 

Coll.  :  Wheeler  468,  Jefferson.     Aug. 
Scutellaria  parvula  MICHX.  Fl.  Bor.  Am.  2:11.     1803. 

Coll.  :   Lyon  39,  Winnebago.     June. 

Agastache  scrophulariaefolia  (WILLD.)  KUNTZE,  Rev.   Gen.  PI. 
511.     1891. 

Coll.:  Wheeler,  465,  Jefferson;   Lyon   315,   Winnebago. 

Aug. 
Nepeta cataria  L.  Sp.  PI.  570.     1753. 

Coll.  :  Lyon  288,  492,  Jefferson.     July. 
Glecoma  hederacea  L.  Sp.  PI.  578.     1753. 

Coll.  :  Lyon  229,  Crooked  Creek.     July. 
Prunella  vulgaris  L.  Sp.  PL  600.     1753. 

Coll.  :  Wheeler  407,  Jefferson.     July. 
Physostegia  virginiana  (L.)  BENTH.    Lab.    Gen.  and  Sp.  504. 

1834- 
Coll.:    Lyon  295,  Jefferson.     July. 

Leonurus  cardiaca  L.  Sp.  PI.  584.     1753. 

Coll.  :  Wheeler  256,  Lyon  418,  Winnebago  ;  Wheeler  295, 
Mayville.     June,  July,  Aug. 

Stachys  palustris  L.  Sp.  PI.  580.     1753. 

Coll.  :  Wheeler  300,  Crooked  Creek.     July. 

Monarda fistulosa  L.  Sp.  PL  22.     1753. 

Coll.  :  Lyon  185,  Crooked  Creek.     July. 
Blephila  hirsuta  (PURSH)  TORR.  FL  U.  S.  27.     1824. 

Coll.:  Lyon   225,    Mayville;  Wheeler   498,   Winnebago. 

July,  Aug. 
Hedeoma  pulegioides  (L.)  PERS.  Syn.  2  :  131.     1807. 

Coll.  :  Wheeler  610,  Brownsville.     Aug. 
Previously  reported  from  the  Mississippi  river  by  Garrison 

and  Miss  Manning,  but  there  are  no  Minnesota  specimens 

in  the  Herbarium  of  the  University. 
Hedeoma  hispida  PURSH,  Fl.  Am.  Sept.  414.     1814. 

Coll.  :  Lyon  51,  Winnebago.     June. 
Clinopodium  vulgare  L.  S  p.  PI.  587.     1753- 

Coll.  :  Lyon  411,  Winnebago.     Aug. 


\\'JlCclcr  :     FLORA.    OF    SOUTHEASTERN    MINNESOTA.  405 

Kcellia  virginiana  (L.)  MAcM.  Met.  Minn.  452.     1892. 

Coll.  :  Wheeler  363,  Crooked  Creek  ;  405,  Jefferson.    July. 
Lycopus  virginicus  L.  Sp.  PI.  21.     1753. 

Coll.  :  Wheeler  537,  Jefferson.     Aug. 
Lycopus  rubellus  MOENCH,  Meth.  Suppl.  146.     1802. 

Coll.  :   Lyon  314,  417,  Winnebago.      Aug. 
Lycopus  americanus  MUHL.  ;  Bart.  Fl.  Phil.  Prodr.  15.     1815. 

Coll.  :  Lyon  335,  348,  Jefferson.     Aug. 
Lycopus  lucidus  TURCZ.  ;  Benth.  in  DC.  Prodr.  12  :  178.      1848. 

Coll.  :   Lyon  459,  Jefferson.     Aug. 
Mentha  canadensis  L.  Sp.  PI.  577.     1753. 

Coll.  :  Lyon  199,  Crooked  Creek.     July. 

SOLANACE^). 

Physalis  philadelphica  LAM.  Encycl.  2:   101.     1786. 

Coll.  :   Lyon  359,  Jefferson.     Aug. 

Mr.  Rydberg  says:  "This  is  a  very  peculiar  form  that  I 
have  never  seen  before.  It  may  be  a  new  species  but  in 
order  to  make  a  good  description  fruit  is  required.  At 
present  it  should  be  referred  to  P.  'philadelphica  with  which 
it  agrees  except  in  the  very  large  and  broad  leaves.  In 
that  respect  it  resembles  P.  macrophysa  Rydb.  but  the 
latter  is  a  perennial  not  an  annual  as  this  plant." 

Physalis  virginiana  MILL.  Gard.  Diet.  Ed.  8,  No.  4.     1768. 
Coll.  :  Lyon  63,  Winnebago.     June. 

Physalis  heterophylla  NEES,  Linnsea,  6:  463.     1831. 
Coll.  :  Lyon  150,  Winnebago.     June. 

Solanum  nigrum  L.  Sp.  PI.  186.     1753. 

Coll.  :  Lyon   192,  Crooked  Creek.     July. 

Datura  tatula  L.  Sp.  PI.  Ed.  2,  256.     1762. 
Coll.  :  Wheeler  379,  Jefferson.     July. 

SCROPHULARIACE^:. 

Verbascum  thapsus  L.  Sp.  PI.  177.     i753- 

Coll.  :  Wheeler  474,  Jefferson.     Aug. 
Scrophularia  marylandica  L.  Sp.  PI.  619.     1753. 

Coll.  :  Lyon  190,  Winnebago  ;  353,  Jefferson.    June,  Aug. 

Chelone  glabra  L.  Sp.  PI.  611.     1753. 

Coll.  :  Wheeler  586,  Crooked  Creek.     Aug. 


406  MINNESOTA    BOTANICAL    STUDIES. 

Mimulus  ringens  L.  Sp.  PL  634.     1753. 

Coll.  :  Wheeler  383,  546,  Jefferson.     July,  Aug. 
Mimulus  jamesii  T.  &  G. ;  BENTH.  in  DC.  Prodr.   10:   371. 

1846. 
Coll.  :    Lyon    68,    Winnebago ;    Wheeler   589,    Crooked 

Creek.     June,  Aug. 
Ilysanthes  gratioloides  (L.)  BENTH.  in  DC.  Prodr.  10:   419. 

1846. 
Coll.:  Lyon  361,  382,  Jefferson  ;  Wheeler  578^,  Crooked 

Creek.     Aug. 
Veronica  americana  SCHWEIN.  ;  BENTH.  in  DC.  Prodr.  10:  468. 

1846. 

Coll.  :  Wheeler  147,  Winnebago.     June. 
Veronica  peregrina  L.  Sp.  PL  14.     1753. 

Coll.:     Lyon  83,    Winnebago;    Wheeler   415,  Jefferson. 

June,  July. 

Leptandra  virginica  (L.)  NUTT.  Gen.  i:   7.     1818. 
Coll.  :  Lyon  234,  250,  Crooked  Creek.     July. 
Dasystoma  grandiflora  (BENTH.)  WOOD.  Bot.  &  Flor.  231.  1873. 

Coll.  :  Wheeler  512,  Winnebago.     Aug. 
This    is    the    first    authentic    specimen    of    this    seen    from 

Minnesota. 
Gerardia  aspera  DOUGL.  :  BENTH.  in  DC.  Prodr.  10:  517.    1846. 

Coll.  :  Lyon  407,  Jefferson.     Aug. 
Gerardia  tenuifolia  VAHL,  Symb.  Bot.  3  :  79.     1794. 

Coll.:  Lyon  406,  456,  Wheeler  575,  Jefferson;    Wheeler 

602,  Crooked  Creek.     Aug. 
Castilleja coccinea  (L.)  SPRENG.  Syst.  2:  775.     1825. 

Coll.  :  Wheeler  85,  Winnebago.    June. 
Castilleja  sessiliflora  PURSH,  Fl.  Am.  Sept.  738.     1814. 

Coll.  :   Lyon  69,  Winnebago.     June. 
Pedicularis  lanceolata  MICHX.  FL  Bor.  Am.  2  :  18.     1803. 

Coll.:  Wheeler  515,  Winnebago;    600,  Crooked   Creek. 

Aug. 

Pedicularis canadensis  L.  Mant.  86.     1767. 
Coll.  :  Lyon  35,  Winnebago.     June. 

LENTIBULARIACE^:. 

Utricularia  vulgaris  L.  Sp.  PL  18.     1753. 

Coll.  ;  Lyon  267,  Wheeler  459,  Jefferson.     July,  Aug. 


Wheeler:   FLORA  OF  SOUTHEASTERN  MINNESOTA.  407 

OROBANCHACE^E. 

Thalesia  uniflora  (L.)    BRITTON,   Mem.   Torr.  Club,  5 :  298. 

1894. 
Coll.  :  Lyon  70,  Winnebago.     June. 

PHRYMACE^E. 

Phryma  leptostachya  L.  Sp.  PI.  601.     1753. 

Coll.  :  Wheeler  298,  Crooked  Creek.     July. 

PLANTAGINACE^E. 

Plantago  major  L.  Sp.  PL  112.     1753. 

Coll.  :  Lyon  124,  Winnebago.     June. 

RUBIACEvE. 

Cephalanthus  occidentalis  L.  Sp.  PL  95.     1753. 

Coll.  :  Wheeler  365,  Crooked  Creek  ;  435,  Jefferson  ;  624, 

Brownsville..    July,  Aug. 

Common  on  the  lowlands  of  the  Mississippi  river. 
Galium  aparine  L.  Sp.  PL  108.     1753. 

Coll.  :  Wheeler  9,  Winnebago.     June. 
Galium  boreale  L.  Sp.  PL  108.     1753. 

Coll.:    Lyon   53,    Winnebago;    199^,   Crooked   Creek. 

June,  July. 
Galium  triflorum  MICHX.  Fl.  Bor.  Am.  i:  80.     1803. 

Coll.  :  Wheeler  41,  Winnebago.     June. 
Galium  trifidum  L.  Sp.  PL  105.     1753. 

Coll.  :  Wheeler  42,  Winnebago.     June. 
Galium  asprellum  MICHX.  Fl.  Bor.  Am.  i :  78.     1803. 

Coll.  :  Wheeler  593,  Crooked  Creek.     Aug. 

CAPRIFOLIACE^E. 

Sambucus  canadensis  L.  Sp.  PL  269.     1753. 

Coll.  :  Wheeler  412,  649,  Jefferson.     July,  Aug. 
Sambucus  pubens  MICHX.  FL  Bor.  Am.  i:   181.     1803. 

Coll.  :  Wheeler  133,  Winnebago.     June. 
Viburnum  opulus  L.  Sp.  PL  268.     1753. 

Coll.:    Lyon    129,   Winnebago;    Wheeler   591,    Crooked 

Creek.     June,  Aug. 
Viburnum  dentatum  L.  Sp.  PL  268.     1753. 

Coll.  :  Wheeler  201,  Winnebago.     June. 


408  MINNESOTA    BOTANICAL    STUDIES. 

Viburnum  lentago  L.  Sp.  PL  268.     1753. 

Coll.  :  Wheeler  39,  Winnebago.     June. 
Triosteum  perfoliatum  L.  Sp.  PL  176.     1753. 

Coll.  :  Wheeler  2,  Winnebago.     June. 
Lonicera  dioica  L.  Syst.  Ed.  12,  165.     1767. 

Coll.  :  Wheeler  190,  Winnebago.     June. 
Lonicera  sullivantii  A.  GRAY,  Proc.  Amer.  Acad.  19  :  76.    1883. 

Coll.  :  Wheeler  122,  Winnebago.     June. 
Diervilla  diervilla   (L.)  MACM.,    Bull.   Torr.   Club,   19:    15. 
1892. 

Coll.  :  Lyon  46,  Winnebago.     June. 

ADOXACE^E. 

AdoxamoschatellinaL.Sp.PL  367.     1753. 

Coll.  :  Wheeler  196,  Winnebago.     June. 
Frequent  in  moist  woods. 

VALERIANACE^:. 

Valeriana  edulis  NUTT.  in  T.  &  G.  Fl.  N.  A.  2  :  48.     1841. 
Coll.  :  Wheeler  159,  Winnebago.     June. 

CAMPANULACE^E. 

Campanula  rotundifolia  L.  Sp.  PL  163.     1753. 

Coll.  :  Lyon  36,  Winnebago.     June. 
Campanula  aparinoides  PURSH,  Fl.  Am.  Sept.  159.     1814. 

Coll.  :  Lyon  194^,  Crooked  Creek.     July. 
Campanula  americana  L.  Sp.  PL  164.     1753. 

Coll.  :  Wheeler  339,  Crooked  Creek.     July. 
Legouzia  perfoliata  (L.)  BRITTON,  Mem.  Torr.  Club,  5 :  309. 
1894. 

Coll.  :  Lyon  148,  Winnebago.     June. 
Lobelia  cardinalis  L.  Sp.  PL  930.     1753. 

Coll.  :  Wheeler  464,  Jefferson  ;  578,  Crooked  Creek.   Aug. 
Lobelia  syphilitica  L.  Sp.  PL  931.     1753. 

Coll.  :  Lyon  310,  340,  Jefferson.     Aug. 
Lobelia  spicata  LAM.  Encycl.  3:  587.     1789. 

Coll.  :  Lyon  115,  Jefferson.     June. 
Lobelia  inflata  L.  Sp.  PL  931.     1753. 

Coll.:  Wheeler  480,   554,  Jefferson;    612,   Brownsville. 
Aug. 


Wheeler  :   FLORA  OF  SOUTHEASTERN  MINNESOTA.          409 

CICHORIACE^E. 

Cichorium  intybus  L.  Sp.  PI.  813.     1753. 

Coll.  :  Wheeler  594,  Lyon  468,  Crooked  Creek.     Aug. 
Adopogon  virginicum  (L.)  KUNTZE,  Rev.  Gen.  PI.  304.     1891. 

Coll.  :    Lyon  20,  337,   Wheeler  506,   Winnebago.     June, 

Aug. 
Taraxacum  taraxacum  (L.)  KARST.  Deutsch.  Fl.  1138.     1880 

-S3- 
Coll.:  Lyon  121,  Winnebago.     June. 

Sonchus  asper  (L.)  ALL.  Fl.  Fed.  i :  222.     1785. 

Coll.  :  Lyon  264,  Jefferson.     July. 
Lactuca  scariola  L.  Sp.  PI.  Ed.  2,  1119.     1763. 

Coll.  :  Lyon  374,  Jefferson.     Aug. 
Lactuca  ludoviciana  (NUTT.)  DC.  Prodr.  7:   141.     1838. 

Coll.  :  Lyon  285,  445,  447,  Jefferson.     July. 
Previously  reported  only  by  Sheldon  from  Sleepy  Eye. 
Lactuca  sagittifolia  ELL.  Bot.  S.  C.  &  Ga.  2:  253.     1821-24, 

Coll.  :  Lyon  363,  Jefferson.     Aug. 

Previously  reported  only  by  Sheldon  from  Lake  Benton. 
Lactuca  floridana  (L.)  GAERTN.  Fruct.  &  Sem.  2  :  362.     1791. 

Coll.  :   Lyon  334,  410,  423,  Winnebago.     Aug. 
Hieracium  umbellatum  L.  Sp.  PI.  804.     1753. 
Coll.  :  Wheeler  627,  Brownsville.     Aug. 
No  authentic  specimens  previously  reported  from  Minnesota. 
Hieracium  canadense  MICHX.  Fl.  Bor.  Am.  2  :  86.     1803. 

Coll.  :  Wheeler  443,  Jefferson;  Lyon  311,  341,  Wheeler 

510,  Winnebago.     Aug. 
Hieracium  scabrum  MICHX.  Fl.  Bor.  Am.  2:  86.     1803. 

Coll.:    Wheeler    485,    Lyon    336,    405,    462,   Jefferson; 
Wheeler  513,  Winnebago;  Wheeler  636,   Brownsville. 
Nabalus  albus  (L.)  HOOK.  Fl.  Bor.  Am.  i:   294.     1833. 
Coll.  :  Wheeler  521,  Winnebago.     Aug. 

AMBROSIACE^E. 
Ambrosia  trifida  L.  Sp.  PL  987.     1753. 

Coll.  :  Wheeler  417,  Jefferson.     July. 
Ambrosia  artemisiaefolia  L.  Sp.  PI.  987.     1753- 

Coll.:   Lyon  495,  Jefferson.     Aug. 
Xanthium  canadense  MILL.  Gard.  Diet.  Ed.  8,  No.  2.    1768. 

Coll.  :  Lyon  424,  Winnebago.     Aug. 


410  MINNESOTA    BOTANICAL    STUDIES. 

COMPOSITE. 

Vernonia  fasciculata  MICHX.  Fl.  Bor.  Am.  2:  94.     1803. 

Coll.  :  Wheeler,  408,  Jefferson.     July. 
Eupatorium  purpureum  L.  Sp.  PI.  838.     1753. 

Coll.:  Lyon  347,  Wheeler  563,  Jefferson ;  Wheeler  519, 

Winnebago.     Aug. 
Eupatorium  altissimum  L.  Sp.  PL  837.     1753. 

Coll.:  Wheeler  533,  568,  Jefferson;  Lyon  412,    Winne- 
bago.    Aug. 
Eupatorium  perfoliatum  L.  Sp.  PI.  838.     1753. 

Coll.  :  Wheeler  493,  Jefferson.     Aug. 
Eupatorium  ageratoides  L.  f.  Suppl.  355.     1781. 

Coll.  :  Lyon  293,  Jefferson.     July. 
Kuhnia  eupatorioides  L.  Sp.  PL  Ed.  2,  1662.     1763. 

Coll.:    Wheeler   532,    555,    571,    Lyon    370,     381,    489, 

Jefferson.     Aug. 

Lacinaria  cylindracea  (Micnx.)  KUNTZE,  Rev.  Gen.  PL  349. 
1891. 

Coll.  :  Lyon  181,  Crooked  Creek;    290,  Jefferson.     July. 
Lacinaria  pycnostachya  (Micnx.)  KUNTZE,  Rev.  Gen.  PL  349 
1891. 

Coll.:  Lyon  265,  Jefferson.     July. 
Lacinaria  scariosa  (L.)  HILL,  Veg.  Syst.  4:  49.     1762  . 

Coll.  :   Lyon  313,  Winnebago.     Aug. 
Solidago  flexicaulis  L.  Sp.  PL  879.     1753. 

Coll.  :  Lyon  371  ^,  484,  Jefferson  ;  Wheeler  590,  Crooked 

Creek.     Aug. 
Solidago  hispida  MUHL.  ;  WILLD.  Sp.  PL  3:  2063.     1804. 

Coll.  :   Lyon  294,  Jefferson.     July. 
Solidago  erecta  PURSH,  FL  Am.  Sept.  542.     1814. 

Coll.  :  Lyon  371,  Jefferson.     Aug. 
Reported  from  Stearns  County  but  no  Minnesota  specimens 

previously  seen. 
Solidago  speciosa  NUTT.  Gen.  2  :   160.     1818. 

Coll.  :  Lyon  467,  Jefferson.     Aug. 
Solidago  ulmifolia  MUHL.  ;    WILLD.  Sp.  PL  3  :   2060.     1804. 

Coll.  :  Lyon  433,  Winnebago.     Aug. 
Solidago  serotina  AIT.  Hort.  Kew.  3:  211.     1789. 

Coll.  :  Lyon  360,  Jefferson.     Aug. 


k  Wheeler :   FLORA  OF  SOUTHEASTERN  MINNESOTA.  411 

Solidago  missouriensis  NUTT.  Journ.  Acad.  Phila.  7:  32.    1834. 
Coll.  :  Lyon  289,  373,  Jefferson.     July. 

Solidago  canadensis  L.  Sp.  PI.  878.     1753. 
Coll.  :   Lyon  330,  Winnebago.     Aug. 

Solidago  nemoralis  AIT.  Hort.  Kew.  3:  213.     1789. 
Coll.:  Wheeler  61 1,  Brownsville.     Aug. 

Solidago  rigida  L.  Sp.  PI.  880.     1753. 

Coll.:  Lyon  372,  403,  Jefferson.     Aug. 

Euthamia  graminifolia  (L.)  NUTT.  Gen.  2:  162.     1818. 
Coll.  :  Wheeler  565,  Jefferson.     Aug. 

Boltonia  asteroides  (L.)  L'HER.  Sert.  Angl.  27.     1788. 
Coll.  :  Lyon  385,  Wheeler  561,  Jefferson.     Aug. 

Aster  drummondii  LINDL.   in  Hook.  Comp.  Bot.  Mag.  i :  97. 

^1835- 

Coll.:    Wheeler  551,  Jefferson;  Lyon  426,  Winnebago. 

Aug. 
Aster  sagittifolius  WILLD.  Sp.  PI.  3:  2035.     1804. 

Coll.:     Lyon    402,   451,   Jefferson;     Wheeler    583,    599, 

Crooked  Creek.     Aug. 
Aster  patens  AIT.  Hort.  Kew.  3:  201.     1789. 

Coll.  :  Wheeler  584,  Crooked  Creek.     Aug. 
Aster  novae-angliae  L.  Sp.  PI.  875.     1753. 

Coll.:  Lyon  425,  Winnebago  ;  482,  Jefferson.     Aug. 
Aster  puniceus  L.  Sp.  PI.  875.     1753. 

Coll.  :  Wheeler  585,  Crooked  Creek.     Aug. 
Aster  prenanthoides  MUHL.  ;  Willd.  Sp.  PL  3  :  2046.     1804. 

Coll.  :  Wheeler  518,  Lyon 409,  Winnebago  ;  Wheeler  582. 

Crooked  Creek.     Aug. 
Aster  laevis  L.  Sp.  PI.  876.     1753. 

»Coll.  :  Wheeler  509,  Lyon  435,  Winnebago  ;  Wheeler  616, 
626,  Brownsville.     Aug. 

Aster  sericeus  VENT.  Hort.  Cels.  pi.  jj.     1800. 
Coll.  :  Wheeler  218,  Jefferson.     June. 

Aster  ptarmicoides  (NEES)  T.  &  G.  Fl.  N.  A.  2  :  160.     1841. 
Coll.:    Lyon  441,  Jefferson.     Aug. 

Aster  salicifolius  LAM.  Encycl.  1 :  306.     1783. 

Coll.:    Wheeler  552,    Lyon   400,  Jefferson;    Lyon  427, 
Winnebago.     Aug. 


412  MINNESOTA    BOTANICAL    STUDIES. 

Aster  paniculatus  LAM.  Encycl.  i  :  306.     1783. 

Coll.:  Wheeler  540,  Jefferson ;   623,  Brownsville.     Aug. 

Erigeron  pulchellus  MICHX.  Fl.  Bor.  Am.  2:   124.     1803. 

Coll.  :  Lyon  28,  Winnebago.     June. 
Erigeron  animus  (L.)  PERS.  Syn.  2  :  431.     1807. 

Coll.:    Lyon  84,   420,    432,  Winnebago;    Wheeler   562, 

Jefferson.     Aug. 
Erigeron  ramosus  (WALT.)  B.S.P.  Prel.  Cat.  N.  Y.  27.     1888. 

Coll.  :  Wheeler  530,  572,  Jefferson.     Aug. 
Leptilon  canadense  (L.)  BRITTON,  in  Brit.  &  Brown,  111.  Fl.  3 : 

391.  f.3827.     1898. 
Coll.  :  Lyon  303,  Jefferson.     Aug. 
Doellingeria  umbellata  pubens  (A.  GRAY)  BRITTON,  in  Brit.  & 

Brown,  111.  Fl.  3:   392.      1898. 
Coll.:     Lyon     399,   Jefferson;    Wheeler     581,    Crooked 

Creek.      Aug. 
Antennaria  plantaginifolia  (L.)  RICHARDS.  App.  Frank.  Journ. 

Ed.  2,'  30.     1823. 

Coll.  :  Lyon  23,  Winnebago.     June. 
Gnaphalium  obtusifolium  L.  Sp.  PI.  851.     1753. 

Coll.  :  Lyon  338,  440,  465,  Jefferson.     Aug. 
Polymnia  canadensis  L.  Sp.  PI.  926.     1753. 

Coll.:    Wheeler    316,    Mayville ;     Lyon    344,  Jefferson. 

July,  Aug. 
Silphium  perfoliatum  L.  Sp.  PL  Ed.  2,  1301.     1763. 

Coll.  :  Lyon  215,  240,  Crooked  Creek.     July. 
Silphium  laciniatum  L.  Sp.  PI.  919.     1753. 

Coll.  :  Wheeler  297,  Crooked  Creek.     July. 
Heliopsis  helianthoides  (L.)  B.S.P.  Prel.  Cat.  N.  Y.  28.     1888. 

Coll.  :  Wheeler  614,  Brownsville.     Aug. 
Heliopsis  scabra  DUNAL,  Mem.  Mus.  Paris,  5  :  56.  pL  4.     1819. 

Coll.  :  Wheeler  330,  Crooked  Creek.     July. 
Rudbeckia  triloba  L.  Sp.  PI.  907.     1753. 

Coll.:   Wheeler  502,  Winnebago  ;  535,  Jefferson.     Aug. 
Not  previously  reported  from  Minnesota.     Infrequent,  edges 

of  thickets. 
Rudbeckia  hirta  L.  Sp.  PI.  907.     1753. 

Coll.:  Wheeler  301,  329,  Crooked  Creek;  613,  Browns- 
ville.    July,  Aug. 


\\'Jiceler :   FLORA  OF  SOUTHEASTERN  MINNESOTA.          413 

Rudbeckia  laciniata  L.  Sp.  PL  906.     1753. 

Coll.  :  Lyon  232,  Crooked  Creek  ;  Wheeler  542,  Jefferson. 

July,  Aug. 
Ratibida  pinnata  (VENT.)   BARNHART,  Bull.  Torr.  Club,  24: 

410.     1897. 
Coll.  :  Wheeler  374,  Crooked  Creek.     July. 

Ratibida  columnaris  (SiMs)  D.  DON  ;  Sweet,  Brit.  Fl.  Card.  2  : 

361.     1838. 

Coll.  :  Wheeler  536,  Jefferson.     Aug. 
Not  previously  reported  from  eastern  Minnesota.     Rare,  dry 

banks. 

Helianthus  atrorubens  L.  Sp.  PI.  906.     1753. 
Coll.  :  Wheeler  634,  Brownsville.     Aug. 
Not  previously  reported  from  Minnesota.     The  only  previous 
collection  known  is  that  of  Sandberg,  Hennepin  Co.,  Aug., 
1889. 

Helianthus  scaberrimus  ELL.  Bot.  S.  C.  &  Ga.  2 :  423.     1824. 
Coll.  :  Wheeler  445,  528,  Jefferson  ;  579,  Crooked  Creek. 

Aug. 
Helianthus  occidentalis  RIDDELL,   Suppl.   Cat.    Ohio  PL    13. 

1836. 
Coll.:  Wheeler  444,   Lyon  322,  Jefferson;  Wheeler  511, 

Winnebago.     Aug. 
Helianthus  grosse-serratus  MARTENS,  Sel.  Sem.  Hort.  Loven. 

1839. 

Coll.  :  Wheeler  549,  Jefferson.     Aug. 
Helianthus  divaricatus  L.  Sp.  PL  906.     1753. 

Coll.  :    Wheeler    566,    576,   Jefferson ;    630    Winnebago. 

Aug. 
Helianthus  tracheliifolius   MILL.  Gard.  Diet.  Ed.   8,   No.   7. 

1768. 
Coll.  :  Wheeler  603,  Crooked  Creek.     Aiig. 

Helianthus  strumosus  L.  Sp.  PL  905.     1753- 
Coll.  :  Wheeler  632,  Brownsville.     Aug. 

Helianthus  tuberosus  L.  Sp.  PL  905.     1753. 
Coll.  :  Wheeler  567,  Jefferson.     Aug. 

Coreopsis  palmata  NUTT.  Gen.  2  :   180.     1818. 

Coll.:     Lyon    160,    Winnebago;     182,    Crooked    Creek. 
June,  July. 


414  MINNESOTA    BOTANICAL    STUDIES. 

Bidens  laevis  (L.)  B.S.P.  Prel.  Cat.  N.  Y.  29.     1888. 

Coll.  :  Wheeler  560,  Jefferson.     Aug. 

Bidens   comosa   (A.  GRAY)  WIEGAND,   Bull.  Torr.    Club,  24: 
^  436.      1897. 

Coll.  :  Wheeler  544,  559,  Jefferson  ;  628,  Brownsville.  Aug. 
Not  previously  reported    from   Minnesota.     The    only    pre- 
viously collected  authentic  specimen  seen  from  Minnesota 

is  that  of  Aiton,  Minneapolis,  Sept.,  1890. 
Common  on  the  low  wet  ground  throughout. 
Bidens  frondosa  L.  Sp.  PL  832.     1753. 

Coll.  :  Wheeler  541,  Jefferson.     Aug. 
Helenium  autumnale  L.  Sp.  PI.  886.     1753. 

Coll.  :  Wheeler  487,  Jefferson.     Aug. 
Achillea  millefolium  L.  Sp.  PL  899.     1753. 

Coll.  :  Wheeler  398,  Jefferson.     July. 
Anthemis  cotula  L.  Sp.  PL  894.     1753. 

Coll.  :  Lyon  269,  284,  Jefferson.     July. 
Chrysanthemum  leucanthemum  L.  Sp.  PL  888.     1753. 

Coll.  :  Lyon  231,  Crooked  Creek.     July. 
Tanacetum  vulgare  L.  Sp.  PL  844.     1753. 

Coll.  :   Lyon  416,  Winnebago.     Aug. 
Artemisia  dracunculoides  PURSH,  Fl.  Am.  Sept.  742.     1814. 

Coll.:    Wheeler  370,    Crooked    Creek;    Lyon  369,   390, 

Jefferson.     Aug. 
Artemisia  serrata  NUTT.  Gen.  2  :  142.     1818. 

Coll.:  Lyon  383, Jefferson.     Aug. 
Artemisia  gnaphalodes  NUTT.  Gen.  2:  143.     1818. 

Coll.  :    Wheeler  550,  Jefferson  ;   Lyon   419,  Winnebago. 

Aug. 
Erechtites  hieracifolia  (L.)  RAF.  DC.  Prodr.  6:  294.     1837. 

Coll.  :   Lyon  342,  446,  Jefferson;  Wheeler  621,  Browns- 
ville.    Aug. 
Mesadenia  reniformis  (MUHL.)  RAF.  New  Fl.  4:  79.     1836. 

Coll.  :  Wheeler  273,  Crooked  Creek.     June. 
Senecio  plattensis  NUTT.  Trans.  Am.  Phil.  Soc.  (II.)  7:  413. 
1841. 

Coll.  :  Wheeler  100,  Winnebago.     June. 
Not  previously  reported  from  Minnesota.     The  only  previous 

known   collection  in  this    state  is  that  of   Prof.    Conway 

MacMillan  from  Hennepin  county. 


Wheeler  :   FLORA  OF  SOUTHEASTERN  MINNESOTA.  415 

Senecio  aureus  L.  Sp.  PI.  870.     1753. 

Coll.  :  Lyon  54,  Winnebago.     June. 
Arctium  minus  SCHK.  Bot.  Handb.  3:  49.     1803. 

Coll.  :  Wheeler  432,  Jefferson.     July. 
Carduus  lanceolatus  L.  Sp.  PI.  821.     1753. 

Coll.:  Lyon  242,  Crooked  Creek.     July. 
Carduus  discolor  (MUHL.)  NUTT.  Gen.  2  :   130.     1818. 

Coll.  :  Lyon  377,  Jefferson.     Aug. 

Carduus   odoratus    (MuHL.)   PORTER.   Mem.   Torr.    Club,    5: 
345.     1894. 

Coll.  :   Herb.  Wheeler  25,  Winnebago.     July. 

DESCRIPTION  OF  PLATE  XXI. 

A.  Juniper  point,  Crooked  creek  valley.     Southwest  side  of  bluff 
dotted  with  junipers  and  white  birch. 

B.  Base  of  bluff,  upper  Winnebago  valley.     White  pine,  juniper 
and  white  birch  along  the  upper  edge  of  cliff. 

PLATE  XXII. 

A.  Western  slope  of  bluff.  The  woods  follow  the  areas  of  greatest 
moisture  i.  «?.,  the  ravines  and  foot  of  bluff  and  the  water  course  in  the 
valley.  The  shrubs  in  the  valley  mark  the  course  of  a  small  creek 
and  are  principally  willows  and  dogwoods. 

£>.  Southern  slope  of  bluff  showing  the  steep  bare  slopes  and  the 
thickly  wooded  ravine.  The  extreme  base  of  the  bluff  to  the  left  has 
been  cleared  of  timber  for  cultivation. 

PLATE  XXIII. 

A.  Grove  of  white  birch. 

B.  Swamp  vegetation.      Spathyema  growing  in  the  shade  of  black 
ash  and  yellow  birch. 

PLATE  XXIV. 

A.  Group  of  coffee  trees  (Gymnocladus). 

B.  White  birch  and  juniper  on  side  of  bluff. 

PLATE  XXV. 

A.  Slough  and  island  vegetation.      Sagittarias  and  Nelumbo  are 
the  most  prominent  water  plants,  and  willows  and  cottonwood  on  the 
island  in  the  background. 

B.  General  view  of  river  valley  from  bluffs  on  Minnesota  side  of 
river.     The  river  channel  is  on  the  farther  side  at  the  base  of  the  Wis- 
consin bluffs. 


416  MINNESOTA    BOTANICAL    STUDIES. 

PLATE  XXVI. 

A.  General  view  of  Winnebago  valley  showing  general  distribution 
of  forest  vegetation.     The  valley  is  almost  entirely  cleared  for  cultiva- 
tion. 

B.  South  branch  of   Winnebago  valley.     The  northern  slope   of 
bluff  is  densely  wooded. 

PLATE  XXVII. 

A.  Lilium  canadense  growing  in  moist  meadow  of  creek  valley. 

B.  Pond    vegetation.     Yellow    pond-lily  with  water  grasses    and 
sedges. 


MINNESOTA  BOTANICAL  STUDIES. 


PART  IV. 


PLATE  XXI. 


VOL.  II. 


MINNESOTA  BOTANICAL  STUDIES. 


PART  IV 


PLATE  XXII. 


THE    MELIOTYPE    PSINTINO    CO.,    BOSTON 


II. 


MINNESOTA  BOTANICAL  STUDIES. 


PART  IV. 


PLATE  XXIII. 


II. 


MINNESOTA  BOTANICAL  STUDIES. 


PART  IV. 


PLATE  XXIV. 


MINNESOTA  BOTANICAL  STUDIES. 


PART  IV. 


'- 

• 

'". 


PLATE  XXV 


THE    HELIOTYPE    PRINTING    CO.,    BOSTON. 


MINNESOTA  BOTANICAL  STUDIES. 


PART  IV. 


PLATE  XXVI. 


THE    HELIOTYPE    PRINTING    CO.,    BOSTON. 


L.  II. 


MINNESOTA  BOTANICAL  STUDIES 


PART  IV 


PLATE  XXVII. 


XXIII.     THE  SEED  AND  SEEDLING  OF  THE 

WESTERN      LARKSPUR     (Delphinium 

occidentale  Wats.). 


FRANCIS  RAMALEY. 


The  seeds  of  Delphinium  occidentale  vary  in  color  from  a 
yellowish  brown  to  a  brownish  black.  The  testa  is  somewhat 
irregularly  roughened  but  not  pitted  or  rugose  as  in  many 
species  of  the  genus,  e.  g.,  the  official  species,  D.  staphisagria. 
The  seeds  are  three  angled  with  rounded  sides  and  bluntly 
pointed  at  the  ends.  The  edges  are  either  merely  sharp  angled 
or  else  the  angles  project,  forming  conspicuous  wings.  (See 
Fig.  4  and  5.)  The  seeds  are  anatropous  as  in  other 
Ranunculaceae.  The  vascular  bundle  extending  from  the 
hilum  is  small,  about  80  microns  in  diameter.  It  is  situated  in 
the  parenchyma  of  one  of  the  angles.  The  cells  of  the  bundle 
are  about  2  or  3  microns  in  diameter,  in  cross  section. 

Endosperm. — The  body  of  the  seed  within  the  seed-coat  con- 
sists chiefly  of  endosperm,  the  embryo  being  very  small.  (See 
Fig.  5  and  6.)  In  the  endosperm,  two  distinct  portions  may 
be  recognized.  The  inner  portion,  an  ellipsoidal  mass,  is  rich 
in  oily  matter.  The  outer  portion  contains  some  oil,  but  the 
cubical  or  prismatic  cells  of  which  it  is  composed  are  chiefly 
filled  with  proteid  grains.  There  is  no  starch  present  in  any 
part  of  the  seed. 

Embryo. — The  embryo,  which  exhibits  slight  differentiation, 
is  placed  at  the  micropylar  end  of  the  seed.  It  is  embedded  in 
the  inner  endosperm.  The  embryo  is  small,  about  0.4  mm. 
long  or  one-fifth  the  length  of  the  entire  seed.  (See  Fig.  5). 

Seed-coat. — The  testa  consists  of  a  large-celled  epidermis 
with  a  thick  cuticle  and  of  four  or  five  layers  of  large-celled  par- 
enchyma. (See  Fig.  14.)  These  cells  have  yellow  or  brownish 
walls  and  contain  only  air.  They  are  usually  very  much  flat- 

417 


418  MINNESOTA    BOTANICAL    STUDIES. 

tened  in  the  dry,  ripe  seed  but  swell  out  in  seeds  which  have 
been  soaked  in  water.  The  tegmen  consists  of  a  single  layer  of 
small  rectangular  cells  with  thick  periclinal  and  thin  anticlinal 
walls.  The  cell  walls  are  of  a  dull  brownish  color  and  the  cav- 
ities are  without  contents. 

Germination  takes  place  in  from  four  to  six  weeks  when 
seeds  are  placed  under  favorable  conditions.  The  cotyledons 
generally  escape  from  the  seed-coat  before  appearing  above 
ground.  This  is  easily  done  because  by  this  time  the  endo- 
sperm has  been  completely  used  and  the  seed-coat  is  likely  to 
be  somewhat  rotted  during  the  long  period  of  germination. 

Morphology  of  the  Seedling. — In  the  young  seedling  the  cot- 
yledons are  small,  the  blades  being  generally  about  4  mm.  in 
length  when  they  first  emerge  above  the  surface  of  the  soil. 
They  increase  considerably  in  size,  becoming  8  mm.  long  and 
6  mm.  wide.  They  are  ovate,  bluntly  pointed,  with  three  prin- 
cipal veins  from  which  spring  conspicuous  secondary  veins. 
The  petioles  are  connate  from  their  bases  to  a  point  only  a  few 
millimeters  from  the  blades.  The  structure  formed  of  the  united 
petioles  emerges  above  the  surface  of  the  soil  in  the  form  of  an 
arch,  thus  simulating  a  hypocotyl.  (See  Fig.  i.)  The  connate 
bases  of  the  cotyledons  form  a  dome-shaped  structure  covering 
the  growing  point  of  the  shoot.  This  structure  may  be  termed 
the  cotyledonary  sheath.  The  development  of  the  foliage  leaves 
causes  a  rupture  of  the  cotyledonary  sheath.  Through  the 
opening  formed  the  first  and  succeeding  leaves  emerge.  (See 
Figs.  2  and  3.)  The  cotyledons  wither  and  finally  disappear 
about  the  sixth  week  after  germination.  The  first  internodes  of 
the  stem  do  not  elongate  and  the  sub-aerial  portion  of  the  plant 
consists  only  of  a  rosette  of  long-petioled  leaves,  until  the  some- 
what scape-like  flowering  stem  is  produced.  The  early  foliage 
leaves  show  considerable  variation  in  the  blade.  The  first  is 
palmately  tri-lobed  with  narrow  sinuses.  In  some  specimens 
the  lobes  are  pointed,  in  others  rounded.  The  separate  lobes 
are  sometimes  rather  deeply  one-  to  two-toothed.  Later  leaves 
may  be  similar  or  may  be  five-lobed,  the  lobes  generally  mucro- 
nate,  or  acute,  not  rounded.  The  young  seedling  of  the  plant 
studied  resembles  that  of  Delphinium  nudicaule,  first  observed 
by  Asa  Gray,*  and  accurately  described  by  Darwin. f  Lub- 

*Graj.     Botanical  Text  Book,  Ed.  VI,  i  :  22.     1879. 

t  Darwin.     The   Power  of  MovementMn  Plants, "p.  80  (American  Edition). 


Ramaley :   SEED  AND  SEEDLING  OF  WESTERN  LARKSPUR.    419 

bock,*  mentions  that  in  Delphinium  troll  if olium  and  in  Del- 
phinium consolida  the  petioles  of  the  cotyledons  are  united  in 
the  same  way. 

Anatomy  of  the  Seedling. — As  this  has  apparently  not  been 
described  for  any  species  of  Delphinium  a  somewhat  extended 
account  will  be  given.  It  may  be  well  to  state  at  the  outset 
that  the  present  writer  has  studied  only  the  seedling  and  not  the 
flowering  stem.  The  young  root  has  a  thick  cortex  and  small 
central  stele.  The  endodermis,  though  thin-walled,  is  con- 
spicuous in  properly  stained  sections  because  of  the  thickened 
cuticularized  spots  on  the  radial  walls.  The  xylem  is  arranged 
in  two  small  groups.  (See  Fig.  7.)  In  an  older  portion  of 
the  root  (Fig.  8)  the  xylem  forms  an  elongated  mass  in  the 
center  of  the  stele.  Higher  up  the  vascular  tissue  extending  to 
the  cotyledons  passes  out  abruptly  on  either  side  at  right  angles 
to  the  longer  diameter  of  the  xylem  mass.  (Fig.  9.)  Passing 
upward  the  xylem  strand  divides  into  six  or  more  bundles  as 
the  transition  occurs  from  root  to  stem.  At  the  same  time 
the  cortex  becomes  thinner.  In  a  cross  section  at  this  point 
(Fig.  10)  the  cotyledonary  sheath  is  seen  surrounding  the  stem. 
In  a  section  somewhat  higher  up  (Fig.  n)  the  bases  of  the  early 
foliage-leaves  may  be  seen  placed  alternately.  Here  the  stem 
abruptly  narrows  and  a  rupture  of  the  cotyledonary  sheath  per- 
mits the  emergence  of  the  first  foliage-leaf.  (Figs.  2  and  3). 
The  cotyledonary  sheath  now  becomes  smaller,  narrowing  to 
form  the  structure  previously  spoken  of  as  resembling  a  hypo- 
cotyl.  Sections  of  this  structure  show  that  its  component 
petioles  are  not  completely  fused  at  any  point  (Fig.  12.)  A 
slit-like  passage,  lined  with  epidermis,  extends  upward  to  the 
point  where  the  petioles  separate  completely. 

Anatomy  of  the  cotyledonary  Sheath. — No  difference  is  to 
be  noted  between  the  outer  epidermis  and  that  lining  the  cavity. 
It  is,  in  both  cases,  composed  of  elongated  cells  which  are 
square  in  cross  section.  There  are  two  vascular  bundles,  one 
for  each  component  petiole.  These  are  small  but  show  no  pe- 
culiarities in  structure.  The  fundamental  tissue  is  a  large- 
celled  parenchyma. 

Anatomy  of  the  Lamina  of  the  Cotyledons.- — Each  lamina 
has  three  principal  veins  which  send  off  numerous  branches. 
The  epidermis  is  composed  of  thin-walled  cells,  somewhat 

*Lubbock.     On  Seedlings,  2:  96.     1892. 


420  MINNESOTA    BOTANICAL    STUDIES. 

larger  on  the  upper  surface  than  on  the  lower.  These  cells 
have  a  very  sinuous  outline  when  seen  in  surface  view.  Sto- 
mata  are  confined  to  the  lower  leaf  surface.  A  loose  palisade 
layer  lies  next  to  the  upper  epidermis.  The  spongy  paren- 
chyma below  this  has  large  air  cavities.  A  few  short,  clavate, 
unicellular  trichomes  sometimes  occur  on  the  under  surface  of 
the  leaf. 

Anatomy  of  the  foliage  leaves. — The  leaves  have  sheath- 
ing bases  and  channeled  petioles.  In  the  center  of  the  petiole 
there  is  an  air  cavity.  Five  or  more  vascular  bundles  form  a 
circle  outside  this  cavity.  (See  Fig.  13.)  Each  bundle  con- 
sists of  a  large  mass  of  xylem,  a  very  small  amount  of  phloem 
and,  external  to  this,  a  small  mass  of  stereom  with  lignified  cell 
walls.  The  fundamental  tissue  is  loose  parenchyma.  No 
special  hypoderma  is  developed.  The  epidermis  is  thin-walled. 
The  leaf  laminae  are  thin  and  composed  of  very  loose  tissue. 
The  epidermal  cells  are  large  and  have  sinuous  outlines.  An 
interesting  peculiarity  is  to  be  noted  in  the  palisade.  The  cells 
of  this  tissue  are  frequently  branched  at  the  upper  end.  (See 
Fig.  15.)  This  peculiarity  was  noted,  according  to  Solereder,* 
by  Haberlandt  in  certain  species  of  allied  genera,  but  that 
investigator  failed  to  find  branched  palisade  cells  in  any  of  the 
species  of  Delphinium  which  he  studied.  The  stomata  of  the 
foliage  leaves  are  confined  to  the  lower  surface  of  the  leaf.  A 
row  of  short,  simple,  pointed  trichomes  is  placed  along  the 
margin  of  the  leaf  and  a  very  few  similar  trichomes  are  scat- 
tered on  the  upper  surface. 

EXPLANATION  OF  FIGURES,  PLATE  XXVIII. 

Figures  i,  2,  3.  Seedlings  of  Delphinium  occidentale  in  various 
stages  of  development  (natural  size).  In  Figures  i  and  2  the  united 
petioles  of  the  cotyledons  have  the  appearance  of  a  hypocotyl.  In 
Figure  2  the  first  leaf  appears  as  a  small  projection  at  the  base  of  the 
petioles  of  the  cotyledons. 

Figure  4.  Seed,      x  18. 

Figure  5.  Longitudinal  section  of  seed  showing  the  minute  embryo. 
The  dotted  ellipse  indicates  the  line  of  division  between  the  inner,  oily 
portion  of  the  endosperm  and  the  outer  part  containing  aleuron  grains. 
X  18. 


*  Solereder.     Sjst.  Anat.  der  Dicotyledonen,  18, 


OL.  II 


MINNESOTA  BOTANICAL  STUDIES. 


PART  IV. 


PLATE  XXV II I. 


Ramaley :   SEED  AND  SEEDLING  OF  WESTERN  LARKSPUR.    421 

Figure  6.  Transverse  section  of  seed  through  the  equator.  The 
vascular  bundle  is  in  the  upper  corner,  x  18. 

Figures  7,  8,  9,  10,  n,  12.  All  x  18.  Diagrams  of  cross  sections 
of  the  seedling  cut  at  various  levels.  Figure  7.  The  root,  thick  cor- 
tex and  small  stele  with  two  xylem  masses.  Figure  8.  The  root, 
higher  up,  a  single  mass  of  xylem.  "  From  this  the  cotyledonary  leaf 
traces  extend  out  horizontally"  (Figure  9).  Figure  10.  The  coty- 
ledonary sheath  with  two  vascular  bundles  encloses  the  stem.  Figure 
1 1 .  The  cotyledonary  sheath  is  ruptured.  The  sheathing  bases  of  the 
foliage  leaves,  arranged  alternately,  enclose  the  small  triangular  apex 
of  the  stem.  Figure  12.  The  united  petioles  of  the  cotyledons  with 
the  slit-like  air  passage. 

Figure  13.  Diagram  of  a  cross  section  of  the  petiole  of  the  first  leaf. 
The  central  air  cavity  is  shown,  also  the  circle  of  vascular  bundles 
(dotted),  each  with  a  small  amount  of  stereom  (black),  x  24. 

Figure  14.  Section  of  a  seed  soaked  in  water.  The  epidermis  has 
a  very  thick  cuticle ;  the  parenchyma  is  large-celled.  The  layer  of 
small  cells  with  thick  walls  is  the  tegmen.  The  endosperm  cells  are 
prismatic  (contents  not  shown).  X  270. 

Figure  15.  Vertical  section  of  the  blade  of  first  foliage  leaf.  Two 
stomata  are  shown  in  the  lower  epidermis.  One  of  the  cells  of  the 
palisade  layer  is  branched  at  the  top.  Chlorophyll  bodies  and  nuclei 
are  shown  in  the  cells.  X  270. 


A    PRELIMINARY    LIST    OF    MINNESOTA 
ERYSIPHE^E. 


E.  M.  FREEMAN. 


The  collection  of  fungi  in  Minnesota  has  been  carried  on  by 
the  Geological  and  Natural  History  Survey  of  the  state  at  various 
times  for  the  past  fourteen  years.*  In  1 886  Prof essorj.  C.  Arthur 
assisted  by  Prof.  L.  H.  Bailey  and  E.  W.  D.  Holway,  Esq.,  made 
a  collection  of  fungi  in  St.  Louis  county  especially  in  the  region 
about  Vermillion  lake.  A  list  of  the  plants  collected  was  pub- 
lished in  Bulletin  No.  3  of  the  Geological  and  Natural  History 
Survey  of  Minnesota.  Since  that  time  numerous  collections 
have  been  made  by  Dr.  A.  P.  Anderson,  Messrs.  E.  P.  Shel- 
don, C.  A.  Ballard  and  others,  but  lists  of  the  collected  plants 
have  not  yet  been  published.  The  list  given  below  comprises 
records  of  all  the  Erysiphese  which  have  been  collected  in  Min- 
nesota up  to  the  present  time  and  deposited  at  the  herbarium  of 
the  University  of  Minnesota. 

A  number  of  specimens  had  been  identified  by  Mr.  Sheldon 
and  these  together  with  the  above  mentioned  collection  of  Profes- 
sor Arthur  and  his  party  have  in  every  case  been  reexamined  so 
that  the  writer  assumes  the  sole  responsibility  of  the  determina- 
tions. The  specimens  have  been  compared  with  such  well 
known  exsiccati  as  Ellis'  North  American  Fungi,  de  Thiimen's 
Mycotheca  Universalis  and  others.  For  the  sake  of  complete- 
ness the  collection  made  by  Professor  Arthur  and  party  is  incor- 
porated in  this  list  and  where  the  nomenclature  has  been  changed 
the  name  published  by  Professor  Arthur  is  placed  in  parenthe- 
ses after  the  collection  citation.  In  18844  A.  B.  Seymour  made 
a  few  collections  along  the  Northern  Pacific  Railroad.  Speci- 

*  A  list  of  Minnesota  fungi  published  by  Dr.  A.  E.  Johnson  in  the  Bulletins  of 
the  Minnesota  Academy  of  Natural  Sciences  during  the  years  1876-1879  cannot  be 
considered  authentic,  since  no  collection  is  available  for  comparison. 

f  Seymour,  A.  B.  List  of  Fungi,  collected  in  1884  along  the  Northern  Pa- 
cific Railroad.  Proc.  Bost.  Soc.  Nat.  Hist.  24 :  182-191.  1889. 

423 


424  MINNESOTA    BOTANICAL    STUDIES. 

mens  of  these  were  not  left  at  the  University  herbarium.  No 
species  however  are  reported  by  him,  that  have  not  been  col- 
lected by  the  staff  of  the  survey.  Mention  of  Seymour's  collec- 
tions is  appended  to  each  species  reported  by  him. 

The  nomenclature  of  Burrill  *  has  been  made  use  of  in  the 
list  and  for  full  synonymy  the  reader  is  referred  to  the  works 
cited  below.  Britton  and  Brown's  Illustrated  Flora  of  the 
United  States  and  Canada  has  been  closely  followed  in  the 
naming  of  all  host  plants. 

Of  the  Erysipheas,  nineteen  species  in  all  have  been  col- 
lected, distributed  among  the  genera  as  follows  :  Spharotheca, 
3  ;  Erysiphe,  5  ;  Uncinula,  3  ;  Phyllactinia,  i  ;  Podosphcera, 
i  ;  Microsphara,  6.  In  field  work  carried  on  during  such  a 
long  period  of  time  and  by  as  many  as  ten  collectors  acting 
independently,  it  is  to  be  expected  that  the  number  of  collec- 
tions of  common  forms  will  be  increased  at  the  expense  of  the 
number  of  species.  A  glance  at  the  list  given  below  will  show 
that  such  has  been  the  case  in  Minnesota.  There  are  undoubt- 
edly at  least  a  dozen  more  species  of  blights  in  the  state,  and 
it  is  hoped  that  this  list  will  aid  future  observations.  In  citing 
the  district  of  collection,  only  the  county  name  is  given. 

1.  Sphaerotheca  humuli   (DC.)  BURRELL,  Bull.  111.  St.  Lab. 

Nat.  Hist.  2  :  400.      1887. 
On  leaves  of : 

Rubus  hispidus  L.  :     St.  Louis,  July,    1886,  Holway  46. 

(JS.  castagnei  Lev.) 

Viola    sp.    indet.  :     Brown,   July,    1891,    Sheldon     851. 
Humulus  lupulus  L.  :  ,f  Sheldon  7020. 

2.  Sphaerotheca  castagnei  LEV.  Ann.  Sci.  Nat.  III.  25:  139. 

1851. 
On  leaves  of : 

Taraxacum  taraxacum  (L.)  KARST.  ;  St.  Louis  (?)  ;  July, 
1886,  Holway  276.      (Not  published  in  Arthur's  report.) 


*  Burrill,  T.  J.,  and  Earle,  F.  S.  Parasitic  Fungi  of  Illinois.  Bull.  111.  St. 
Lab.  Nat.  Hist.  2.  1887. 

Ellis,  J.  B.,  and  Everhart,  B.  M.  North  American  Pyrenomycetes,  2-30. 
1892. 

fMr.  Sheldon's  last  field  note  book  has  not,  up  to  the  present  time,  been 
found.  Consequently  the  dates  of  collection,  the  district  and  the  name  of  the 
host  plant  often  cannot  be  determined.  The  missing  data  are  indicated  as 
above. 


Freeman  :    PRELIMINARY  LIST  OF  MINNESOTA  ERYSIPHE^E.    425 

Pedicularis  lanceolata  'MiCHX.  ;  Lincoln,    August,   1891, 

Sheldon  1522. 

Bidens  frondosa  L.  ;  Lincoln,  August,  1891,  Sheldon 
1516,-  -  September,  1893,  Sheldon  6092,  Hennepin, 
Oct.,  1898,  Freeman  50. 

Seymour  reports  this  species  on  Erechtites  hicracifolia  and 
Nabalus  sp.  at  Lake  Minnetonka. 

3.  Sphasrotheca  mors-uvae  (Scnw.)  B.  &  C.    Grev.  4:  158. 
1876. 

On  leaves  of : 

Ribes floridum  L'HER.  :  St.  Louis,  July,  188,  Holway  84 
(Splicer  othcca  -pannosa  LEV.);  Kandiyohi,  July,  1892, 
Frost  249. 

4.  Erysiphe   cichoracearum  DC.  Flore  Franc.  2:274.     1815. 
On  leaves  of : 

Ambrosia  artemisicefolia  L.  :  Hennepin,  1890,  MacMillan  ; 

,  Sheldon  7322  ;  ,  Sheldon  6162  ; ,  Sheldon 

6131  ;  Ramsey,  Sept.,  1898,  Freeman  61. 

Ambrosia  -psilostachya  DC.  Traverse,  Sept.,  1893,  Sheldon 
7081. 

Ambrosia  trifida  L.  :  Pope,  Aug.,  1891,  Taylor  1126; 
Brown,  Sept.,  1891;  Sheldon  1243;  Traverse,  Sept., 
1893,  Sheldon  7086;  Goodhue,  Aug.,  1893,  Anderson 
727;  Ramsey,  Sept.,  1898,  Freeman  62;  Hennepin, 
Sept.,  1898,  MacMillan. 

Heliopsis  scabra  DUNAL.  :  Kandiyohi,  Aug.,  1892,  Frost 
449. 

Cnicus&p.  indet.  :  Hennepin,  1890,  MacMillan  ;  Traverse, 
Sept.,  1893,  Sheldon  7072. 

Carduus  sp.  indet.  :  ,  Sheldon  7357  ;  Hennepin,  1890* 

MacMillan. 

Aster  puniceus  L.  var.  lucidulus  A.  GRAY  :  Lincoln,  Aug., 
1891,  Sheldon  1507. 

Aster  sp.  indet.:  Winona,  Sept.,  1888,  Holzinger;  Hen- 
nepin, Oct.,  1898,  Freeman  63  ;  Hennepin,  Oct.,  1892, 
Sheldon  4123. 

Solidago  canadensis  L.  :  ,  Sheldon  6082. 

Solidago  sp.  indet.  :  Hennepin,  1890,  MacMillan  ;  Waseca, 
June,  1891,  Taylor  188  ;  Hennepin,  Oct.,  1891,  Sheldon  ; 
Goodhue,  Aug.,  1893,  Anderson  814. 


426  MINNESOTA    BOTANICAL    STUDIES. 

Helianthus   divaricatus    L.  :   ,   Sept.,    1893,  Sheldon 

6089. 

Helianthus  decapetahis  L.  :  Brown,  July,  1891,  Sheldon, 
1244. 

Helianthus  grosse-serratus  MARTINS  :  Traverse,  Sept., 
1893,  Sheldon  7106. 

Helianthus  scaberr.imus  ELL.  :  ,  Sept.,  1893,  Sheldon 

6143  ;  Ramsey,  Sept.,  1898,  Freeman  64. 

Helianthus  tuberostis  L.  :  Goodhue,  Aug.,  1893,  Anderson 
705  ;  Ramsey,  Sept.,  1898,  Freeman  65. 

Heilanthus  sp.  indet.  :  Traverse,  Sept.,  1893,  Sheldon 
7085  ;  Hennepin,  1890,  MacMillan ;  Blue  Earth,  June, 
1891,  Sheldon  483;  Winona,  Sept.,  1888,  Holzinger; 
Lincoln,  Aug.,  1891,  Sheldon  1418. 

Verbena stricta  VENT.  :  Ramsey,  Sept.,  1898,  Freeman  66. 

Verbena  urticifolia  L.  :  Chisago,  Aug.,  1892,  Taylor 
1639;  Hennepin,  Aug.,  1890,  MacMillan;  Hennepin, 
Sept.,  1898,  Freeman  67. 

Verbena  hastataL.  :  Hennepin,  Oct.,  1892,  Sheldon  4128; 
Pope,  Aug.,  1891,  Taylor  1188;  Goodhue,  Aug.,  1893, 
Anderson  827  ;  Ramsey,  Sept.,  1898,  Freeman  68. 

Verbena  sp.  indet.  :  Traverse,  Sept.,  1893,  Sheldon  7030; 
Carver,  July,  i89i,Ballard  650;  Traverse,  Sept.,  1893, 
Sheldon  7071. 

Rudbeckia  laciniata  L.  :  Hennepin,  Sept.,  1898,  MacMil- 
lan. 

Senecio  aureus  L.  :  Brown,  July,  1891,  Sheldon  1153. 

Hydrophyllum  virginicum  L.  :  Blue  Earth,  June,  1891, 
Sheldon  203. 

Grindelia  squarrosa  (PURSH)  DUNAL  :  Pipestone,  Aug., 
1891,  Sheldon  1434. 

Lappula  virginiana  (L.)  GREENE  :  Blue  Earth,  June, 
1891,  Sheldon  483  ;  St.  Louis,  July,  1886,  Holway  78. 

Lappula  sp.  indet.  :  Brown,  Aug.,  1891,  Sheldon  1232. 

Coreopsis  palmata  NUTT.  :   Winona,  Sept.,   1888,   Holz- 
inger. 
5.  Erysiphe  communis  (WALLR.)   FR.  Summa.  Veg.  Scand. 

406.      1849. 
On  leaves  of : 

JSupatorium  ageratoides  L.  :  Hennepin,  Oct.,  1893,  Shel- 
don 4083. 


Freeman :   PRELIMINARY  LIST  OF  MINNESOTA  ERYSIPHE^E.    427 

Lathyrus  venosus  MUHL.  :  Winona,  Sept.,  1888,  Holzinger 
326;  Pope,  July,  1891,  Taylor  1181  ;  Mille  Lacs,  July, 

1892,  Sheldon   2755;     Otter  Tail,  Aug.,  1892,  Sheldon 
3661. 

Lathy  rus  sp.  indet.  ,  Sheldon  6127. 

(Enothera  sp.  indet.  :   Hennepin,  1890,  MacMillan. 

Clematis  virginiana  L.  :   Hennepin,  1890,  MacMillan. 

Anogra  albicaulis  (PuRSH)  BRITTON.  :.  Brown,  July, 
1891,  Sheldon  1195. 

Strophostyles  helvola  (L.)  BRITTON:  Pope,  July,  1891, 
Taylor  1136. 

Falcata  comosa  (L.)  KUNTZE  :  Pope,  July,  1891,  Taylor 
1136. 

Oxygraphis  cymbalaria  ( PURSH)  PRANTL  :  Lincoln, 
Aug.,  1891,  Sheldon  1357. 

Astragalus  canadensis  L.  :  Lincoln,  Aug.,  1891,  Sheldon 
1423;  Hennepin,  Sept.,  1898,  MacMillan. 

Aragallus  involutus  A.  Nels.  :  Lincoln,  Aug.,  1891,  Shel- 
don 1390. 

An   undetermined  plant  of  pea  family:  Traverse,  Sept., 

1893,  Sheldon  7257. 

Thalictrum  purpurascens  L.  :  Chisago,  Sept.,  1893,  Shel- 
don 6188. 

Thalictrum  sp.  indet.  :  Cass.  Aug.,   1893,  Anderson  706. 

Onagra  biennis  (L.)  SCOP.  ,  Sept.,  1893,  Sheldon 

6146. 

Anemone  virginana  L.  :  Traverse,  Sept.,  1893,  Sheldon 
7089. 

Lotus  americanus  (NuTT.)  BISH.  :  Big  Stone,  Sept., 
1893,  Sheldon,  Traverse,  Sept.,  1893,  Sheldon  7201. 

Polygonum  aviculare  L.  :  Ramsey,  Sept.,  1898,  Freeman 

SI' 
Seymour  reports  E.  communis  on  Lathyrusf  at  Lake  Min- 

netonka. 

6.  Erysiphe  aggregate.  (PECK)  FARLOW,  Bull.  Bussey,  Inst.  2  : 

227.     1878. 
On  leaves  of : 
Alnus  incana  (L.)  WILLD.  :  St  Louis,  July,  1886,  Holway 


428  MINNESOTA    BOTANICAL    STUDIES. 

7.  Erysiphe  galeopsidis  DC.  Flore  Franc.  6  :   108.     1815. 
On  leaves  of : 

S tacky s  -palustrts  L.  :  Lincoln,  Aug.,  1891,  Sheldon 
1572;  Lincoln,  Aug.,  1891.  Sheldon  1261. 

Seymour  reports  this  species  on  Stachys  -palustris  at  De- 
troit, Minnesota. 

8.  Erysiphe  graminis  DC.  Flore  Franc.  6  :   106.     1815. 
On  leaves  of: 

Poa  pratensis  L.  :  Waseca,  June,  1891,  Taylor  228.  Xo 
perithecia  found.  Conidial  stage  ( Oidmm  monthoides 
Link)  only  is  present. 

9.  Uncinula  salicis  (DC.)  WINT.  Die  Pilze  i2:  40.     1887. 
On  leaves  of : 

Salix  sp.  indet.  :  Winona,  Sept.,  1888,  Holzinger ;  Chi- 
sago,  Sept.,  1891,  Sheldon  4263  ;  Traverse,  Sept.,  1892, 
Sheldon  7172;  Traverse,  Sept.,  1892,  Sheldon  7013; 
Chisago,  Aug.,  1892,  Taylor  1634;  Otter  Tail,  July, 

1892,  Sheldon   3936;    Hennepin,    Oct.,   1893,  Sheldon 
4093  ;  McLeod,  Aug.,  i.  J.  McElligott ;  Traverse,  Sept., 

1893,  Sheldon  7069;  Ramsey,  Sept.,  1898,  Freeman  60. 
Salix  bebbiana  SARG.  :  Chisago,  Sept.,  1891,  Sheldon  4246. 
Salix  myrtilloides   L.  :    Otter    Tail,  July,    1892,  Sheldon 

3573^- 

Salix  discolor  MUHL.  :  Hennepin,  Oct.,  1893,  Sheldon 
4089. 

Populus  deltoides  MARSH.  :  Hennepin,  July,  1889,  Shel- 
don;  Hennepin,  Oct.,  1889,  MacMillan ;  Wabasha, 
Sept.,  1893,  Edna  Porter. 

Populus  grandidentata  MICHX.  :  Hennepin,  1890,  Mac- 
Millan. 

Populus  tremuloides  MICHX.  :  Goodhue,  Aug.,  1893,  An- 
derson 707. 

Populus  sp.  indet.  Hennepin,  Oct.,  1889,  MacMillan. 

Seymour  reports  U.  salicis  on  Salix  sp.  at  Lake  Minne- 
tonka. 

10.  Uncinula   clintonii  PECK,    Rep.    N.  Y.  St.  Mus.  25 :  96. 

1873.     Trans.  Alb.  Inst.  7:  216. 
On  leaves  of : 

Tilia  americana  L.  :  Winona,  Sept.,  1888,  Holzinger. 


Freeman  :    PRELIMINARY  LIST  OF  MINNESOTA  ERYSIPHE^E.    429 

11.  Uncinula    necator   (ScH\v.)   BURRILL.  N.   A.    Pyren.    15. 

1892. 
On  leaves  of : 

Parthenocissus  quinqiiefolia  (L.)  PLANCH.  :  Ramsey, 
Sept.,  1898,  Freeman  59;  Hennepin,  Sept.,  1898,  F. 
K.  Butters. 

12.  Phyllactinia  suffulta  (REB.)  SACC.  Mich.  2  :  50.     1880. 
On  leaves  of  : 

Tilia  sp.  indet.  :  Le  Sueur,  June,  1891,  Sheldon  64.  Sey- 
mour reports  this  species  on  Betula  ?  at  Lake  Minnetonka. 

13.  Podosphasra  oxyacanthae  (DC.)  D.  BY.  Beitr.  Morph.  und 
Phys.   der  Pilze,  Part  3,  48.      1870. 

On  leaves  of : 

Prunus  sp.  indet.  :   Hennepin,  1890,  MacMillan. 
Crat&gus  (?)  sp.  indet.:    Le   Sueur,  June,  1891,    Sheldon 
62. ;  Wabasha,  Sept.,  1893,  Edna  Porter. 

14.  Microsphaera  russellii   CLINTON,    Rep.    N.    Y.    St.    Mus. 
26  :  80.      1874. 

On  leaves  of : 

Oxalis  stricta  L.  :  Winona,  Aug.,  1888,  Holzinger. 

15.  Microsphaera  ravenelii  BERK.  Grev.  4:  160.     1876. 
On  leaves  of : 

Lathyrus&p.  indet.  :  Goodhue,  Aug.,  1892,  Ballard  1152; 

Goodhue,  Aug.,  1893,  Anderson  708. 
Seymour   reports    M.    ravenelii  Berk,    on    Lathyrus  ?  at 

Detroit,  Minn. 

16.  Microsphaera  quercina  (Scnw.)   BURRILL,     Bull.    111.   St. 
Lab.  Nat.  Hist.  2  :  424.      1887. 

On  leaves  of : 

^jiercus  macrocarpa  MICHX.  :  Hennepin,  1890,  Mac- 
Millan;  Ramsey,  Sept.,  1898,  Freeman  52. 

17.  Microsphaera  symphoricarpi  HOWE,  Bull.  Torr.  Club,  5:  3. 

1874. 
On  leaves  of : 

Symphortcarpos  sp.  indet.:   Hennepin,  1890,  MacMillan; 

Waseca,  June,  1891,  Taylor6i5  ;  Goodhue,  Aug.,  1893, 

Anderson  815  ; ,  Sheldon  7262  ; ,  Sheldon  7392. 

Symphortcarpos  racemosa  MICHX.  :  Traverse,  Sept.,  1893, 

Sheldon  7084. 
Symphortcarpos   symphoricarpos    (L.)   MAcM.  Traverse, 

Sept.,  1893,  Sheldon  7083. 


430  MINNESOTA    BOTANICAL    STUDIES. 

Symphoricarpos  occidentals  HOOK.  :  Ramsey,  Sept.,  1898, 
Freeman  53. 

18.  Microsphaera  diffusa   C.  &  P.   Journ.    of   Bot.    n :    1872. 

Rep.  N.  Y.  St.  Mus.  25:  95.     1873. 
On  leaves  of : 

Lespedeza   violacea    (L.)    PERS.  :    Winona,    Sept.,   1889, 

Holzinger. 

Lathyrus  sp.  indet.  :   Hennepin,  1890,  MacMillan. 
Meibomia  canadensis  (L.)  KUNTZE.  :  Lincoln,  Aug.,  1891, 

Sheldon  1521  ;  ,  Sept.,  1893,  Sheldon  6105. 

Seymour    reports  M.   diffusa    on  Lespedeza    capitata    at 

Brainerd. 

19.  Microsphaera  alni  (DC.)  WINT.  Die  Pilze  i2:38.     1887. 
On  leaves  of : 

Lonicera  sp.  indet.  :  St.  Louis,  July,  1886,  Holway  242, 
(M.  dttbyiLev.)  ;  Ramsey,  Sept.,  1898,  Freeman  56. 

Lonicera  hirsuta  Eaton  :  St.  Louis,  July,  1886,  Holway 
150,  (M.  dubyi  Lev.). 

Syringa  vulgaris  L.  :  Hennepin,  July,  1889,  Sheldon; 

Hennepin,  Oct.,  1891;  ,  Sheldon  5806;  Goodhue, 

Aug.,  1893,  Anderson  714. 

Alnus  sp.  indet.  :  Hennepin,  1890,  MacMillan. 

Viburnum  lentago  L.  :  Waseca,  June,  1891,  Sheldon, 
506^;  Case,  Aug.,  1893,  Anderson  668;  Ramsey, 
Sept.,  1898,  Freeman  54. 

Viburnum  sp.   indet.  :  Ramsey,  Sept.,  1898,  Freeman  55. 

Lonicera  dioica  L.  :  Goodhue,  Aug.,  1893,  Anderson  753. 

Corylus  americana  WALT.  :  Ramsey,  Sept.,  1898,  Free- 
man 57. 

Tilia  americana  L.  !  :  Hennepin,  Oct.,  1898,  Freeman  58. 

Seymour  reports  this  species  on  Ceanothus  americanus  at 
Brainerd  and  on  Syringa  vulgaris  and  Betula  at  Lake 
Minnetonka. 


NATIVE   AND   GARDEN   DELPHINIUMS    OF 
NORTH  AMERICA. 


K.  C.  DAVIS. 


The  name  Delphinium  (Linn.  Sp.  PI.  530,  1753)  is  from  the 
Greek  delphin^  a  dolphin,  from  the  resemblance  of  the  flower, 
The  common  name  is  LARK  SPUR. 

It  is  a  genus  of  beautiful,  hardy  plants,  annual  or  perennial, 
erect,  branching  herbs.  Leaves  palmately  lobed  or  divided ; 
large,  irregular,  showy  flowers  in  a  raceme  or  panicle ;  sepals 
petal-like,  five,  the  posterior  one  prolonged  into  a  spur ;  petals 
two  or  four,  small,  the  two  posterior  ones  usually  spurred,  the 
lateral  or  lower  ones  small  if  present ;  the  few  carpels  always 
sessile,  forming  many-seeded  follicles. 

There  are  probably  more  than  200  species.  In  fact  Huth's 
last  complete  monograph  recognized  198  species  besides  a  num- 
ber of  doubtful  ones.  The  following  treatment  includes  the 
native  and  cultivated  Delphiniums  of  North  America,  52 
species  and  many  varieties  and  garden  forms.  Thirty  species 
are  native  of  America  north  of  Mexico,  thirteen  of  which  are 
used  in  gardens.  Thirteen  Old  World  species  have  been  intro- 
duced int6  the  American  trade.  Nine  Mexican  species  are  dis- 
tinct, and  none  of  them  are  in  use.  The  mark  (f)  after  a  de- 
scription indicates  which  plants  are  not  used  in  the  trade. 
Four  species  are  of  much  greater  popularity  than  the  others  : 
the  annual  D.  Ajacis,  and  the  perennials  D.  grandiflorum,  D. 
hybridum,  and  D.  formosum.  The  last  three  have  been  es- 
specially  prolific  in  giving  us  new  garden  forms. 

In  presenting  this  paper  I  wish  to  extend  thanks  to  those 
who  have  materially  helped  me,  especially  to  those  who  have 
freely  given  me  the  privilege  of  examining  numerous  speci- 
mens :  Dr.  J.  N.  Rose,  Professor  E.  L.  Greene,  Dr.  N.  L. 
Britton  and  Dr.  B.  L.  Robinson. 

431 


432  MINNESOTA    BOTANICAL    STUDIES. 

The  recent  extended  articles  on  the  genus  are  :  A.  Gray, 
"  An  attempt  to  Distinguish  Between  the  American  Delphin- 
iums," Bot.  Gaz.  12 :  49-54,  1887  ;  and  Syn.  Fl.  I  :  45-52, 
1895.  E.  Huth,  "  Monog.  Gattung  Delphinium,"  in  Engl. 
Bot.  Jahrb.  20:  322-499,  1895.  K.  C.  Davis,  in  Bailey's 
Cyclopedia  of  American  Horticulture. 

SYNOPSIS  OF  SPECIES  OF  DELPHINIUM. 

A.   Roots  annual ;   petals  only  2,  united;   follicles  i. 

B.   Follicles  pubescent,  y2  to  \y2  inches  long ..i.  Ajacis. 

BB.   Follicles  glabrous,  ^  to  y2  inch  long 2.consolida. 

A  A.  Roots  perennial ;  petals  4;   follicles  3  to  5. 
B.   Sepals  red. 

C.   Plant  glabrous ;   seeds  smooth 3.  nudicaule. 

CC.  Plant  partly  pubescent ;   seeds  thin  winged 4.  cardinale. 

BB.   Sepals  greenish   yellow,  yellow,  or  sometimes  marked  with 
blue. 
C.   Inflorescence  and  leaves  densely  hairy. 

D.  Flowers  not  tinged  with  blue 5.  viridescens. 

DD.  Flowers  sordid  white  tinged  with  blue... 6.   Calif ornicum. 
CC.   Inflorescence  and  leaves  glabrescent  or  soon  becoming  so. 

D.  Mature  follicles  densely  hairy 7.  Przeivalskii. 

DD.  Mature  follicles  smooth  or  sulcate. 

E.   Seeds  with  plates  or  scales  in  transverse  rows... 8.  Zalil. 

EE.   Seeds  winged  and  somewhat  wrinkled... 9.  viride. 

BBB.  Sepals  blue,  or  varying  to  white,  or  white. 

C.   Species    native    north    of   Mexico,   or    introduced    from    Old 
World. 
D.  Height  \y2  feet  or  less. 

E.  Natives  of  America  north  of  Mexico. 

F.  Petioles  dilating  and  somewhat  sheathing  at  the  base. 
G.   Stem  lax ;  follicles  glabrous  or  becoming  so. 

H.  Roots  fascicled  and  thickened  but  not  tuberiform. 

10.  bicolor. 
HH.   Roots  fasciculately  tuberous,  or  grumose. 

n.  decorum. 

GG.   Stem  rather  stout,  erect :  follicles  pubescent. 
H.  Length  of  sepals  about  equalling  the  petals. 

I.  Seeds  winged  at  the  angles 12.  hesperium. 

II.  Seeds  scaly  and  bur-like 13.  Hanseni. 

HH.   Length  of  sepals  much  gi-eater  than  petals. 

14.  variegatum. 
FF.  Petioles  hardly  dilating  at  the  base. 


Da-vis :   DELPHINIUMS  OF  NORTH  AMERICA.  433 

G.   Coats  of  seeds  smooth ;   roots  fasciculately  tuberous. 

15-  tricorne. 

GG.   Coats  of  seeds  winged  or  wrinkled,  roots  not  tuber- 
ous, but  in  some  grumose. 
H.   Roots  not  grumose. 

I.  Sepals  shorter  than  the  spur. 

J.   Leaves  thickish  ;  racemes  long. 

1 6.    Andersonii. 

JJ.   Leaves  not  thick ;   racemes  shorter ;    flowers 
smaller 17.   Parishii. 

II.  Sepals  as  long  as  spur,  much  surpassing  petals. 

18.   Parry 7. 

HH.   Roots   coarsely  granular   or  grumose;    carpels 
always  3,  seeds  wing-margined. 

I.  Pedicels  longer  than  the  flowers ;  follicles  spread- 

ing when  mature 19.  Menziesii. 

II.  Pedicels  shorter  than  flowers;   follicles  spread- 

ing only  at  tips 20.  pauciflorum. 

EE.  Natives  of  Asia  but  introduced  to  American  gardens. 
F.  Sepals  somewhat  persistent ;  bractlets  opposite,  lanceo- 
late, entire,  near  the  flower 21.  Brunonianum. 

FF.  Sepals  deciduous  ;  bractlets  alternate,  linear,  or  linear- 

lobed,  distant  from  flower 22.    Cashmirianum. 

DD.   Height  more  than  \yz  feet  (except  in  a  few  cases). 
E.  Seeds  wrinkled  or  scaly,  hardly  winged  (except  in  28  and 

29)  ;  all  native  of  the  United  States  except  23. 
F.  Follicles  always  3. 

G.  Upper  petals  violet 23.   altissimum. 

GG.  Upper  petals  yellowish,  or  yellow  with  blue  tips. 
H.  Inflorescence  a  crowded,  erect,  pyramidal  raceme. 

24.   exaltatum. 
HH.  Inflorescence    open  and    somewhat  branching; 

pedicels  long  and  slender 25 .    Treleasi. 

FF.  Follicles  commonly  varying  from  3  to  5  on  each  plant. 
G.  Stems  more  or  less  leafy. 
H.   Sepals  and  spurs  blue. 

I.  Stem  leafy ;  radical  leaves  few. 

26.    Carolinianum. 

II.  Stem  leaves  mostly  near  the  base ;  radical  leaves 

many. 

J.   Root  a  flattish  tuber 27.    Oreganum. 

JJ.  Root  woody-fibrous 28.    Geyeri. 

HH.   Sepals  and  spurs  chiefly  white.. 29.  camporum. 
GG.   Stem  leafless. 


434  MINNESOTA    BOTANICAL    STUDIES. 

H.   Petioles  of  root  leaves  much  longer  than  blades. 

30.  scaposum. 
HH.  Petioles  of  root  leaves  nearly  equalling  blades. 

31.   uliginosum. 
EE.   Seeds  decidedly  winged. 

F.  Upper  petals  white,  never  yellow 32.  trolliifolium. 

FF.  Upper  petals  often  yellow  or  yellowish. 

G.   Species  from  Old  World   introduced  into  gardens ; 

follicles  always  3. 

H.  Lower  petals  deep  blue,  2-lobed,  yellow-bearded. 

33.  elatum. 

HH.  Lower  petals  bright  blue,    entire,   undulate  01 
slightly  2-lobed. 

I.  Flowers  very  large;   spurs  9  to  10  lines  long. 

34.  grandiflorum, 

II.  Flowers  smaller;   spurs  5  to  8  lines  long. 

35.   cheilanthum . 
GG.   Species  from  west  of  the  Rockies  ;  follicles  always 

3- 
H.  Plant  glabrous,  at  least  in  lower  part. 

I.  Roots  fascicled,  not  tuberous  nor  grumose. 

J.  Follicles  pubescent 36.  scoptdorum. 

JJ.  Follicles  glabrous 37.  glaucum. 

II.  Root  tuberous  or  grumose. 

J.   Lower  pedicels  rather  spreading,  longer  than 

the  spurs. 
K.   Sepals  equal  to  spur  in  length. 

38.  glaucescens. 
KK.   Sepals  shorter  than  the  spur. 

39.  Nuttallii. 
JJ.   Lower  pedicels  and  others  appressed,  shorter 

than  spurs 40.  distichum. 

HH.  Plant  pubescent  throughout 41.  simplex. 

EEE.  Seeds  scaly;  lower  petal  2-lobed;  Old  World  type. 
F.  Petioles  hardly  dilating  at  base,  not  sheathing ;  lower 

petals  yellow  bearded  42.  formosum. 

FF.  Petioles  sheathing  at  base ;  beard  on  lower  petals  not 
yellow. 

G.  Flowers  in  loose  panicles 43.   Maackianum. 

GG.  Flowers  in  dense  racemes 44.  hybridum. 

CC.  Natives  of  Mexico,  not  introduced  to  American  gardens. 
D.  Carpels  puberulent  to  hairy  at  first. 

E.  Plant  glandular-hispid  above 45.  Madrense. 

EE.  Plant  not  glandular-hispid  in  upper  parts. 


Da-cis:   DELPHINIUMS  OF  NORTH  AMERICA.  435 

F.   Lower  petals  provided  with  a  scale-like  appendage  at 
base. 

G.   Stamens  puberulent 46.   bicornutum. 

GG.   Stamens  glabrous. 

PI.  Upper  petals  tipped  with  yellow.. 47.  Ehrenbergi. 
HH.  Upper  petals  not  tipped  with  yellow. 

48.  pedatisectum. 

FF.   Lower  petals  with  appendages  wanting  or  obscure. 
G.   Leaves  pubescent  or  villose. 

H.  Upper  petals  blue 49.  latisepalum. 

HH.   Upper  petals  yellow  with  blue  tips. 

50.   tenuisectum. 

GG.   Leaves  glabrous 51.  leptophyllum. 

DD .   Carpels  glabrous 52.    Wislizeni. 

1.  D.  Ajacis  LINN.  Sp.  PL  531.     1753. 

D.    consolida    SIBTH.   &   SM.    Fl.   Gr^eca,  Prod.   1 :    370. 

1806.     Not  L. 

D.  ornatum  BOUCHE,  in  Bot.  Zeit.  1 :  26.     1843. 
D.-pubescens  GRISEB.  Spicil.  Fl.  Rumel.  i:  319.     1843. 

Not  DC. 
Ceratosanthus     a/act's     SCHUR.     Enum.    PI.    Transs.    30. 

1866. 
f  D.  addendum  McNAB.  in  Trans.  Bot.  S.  Edinb.  9:  335. 

1868. 

An  erect  annual  about  18  inches  high  with  a  few  spreading 
branches  :  leaves  of  stem  sessile,  deeply  cut  into  fine  linear 
segments ;  root-leaves  similar  but  short-petioled :  flowers 
showy,  blue  or  violet,  varying  to  white,  more  numerous  than 
in  D.  consolida^  in  a  spicate  raceme;  petals  2,  united; 
calyx-spur  about  equalling  the  rest  of  the  flower :  but  one  fol- 
licle, pubescent;  seeds  with  wrinkled,  broken  ridges.  May  to 
Aug.  Europe.  Fl.  Grasca,  t.  540.  Rev.  Hort.  1893,  p.  228. 

2.  D.  consolida  LINN.  Sp.  PI.  530.     1753. 
D.  segctum  LAM.  Fl.  Fr.  3:  325.     1778. 

D.  monophyllum  GILIB.  Fl.  Lithuan.  2:  287.     1781. 
D.  versicolor  SALISB.  Prod.  375.     1796- 
CeratosantJnis  consolida  SCHUR.  Verh.  Sieb.  Ver.  Naturf. 
46.     1853. 

An  erect,  hairy  annual,  i  to  i^  feet  high:  leaves  similar  to 
D.  Ajacis:  flowers  few,  loosely  panicled,  pedicels  shorter  than 


436  MINNESOTA    BOTANICAL    STUDIES. 

the  bracts,  blue  or  violet  or  white;  petals  2,  united:  follicle  i, 
glabrous;  seeds  with  broken,  transverse  ridges.  June  to  Aug. 
Europe.  Baxter  Brit.  Bot.  4.  t.  297.  Rev.  Hort.  1893,  p.  228 
(var.  ornatum  candelabrum}. 

3.  D.  nudicaule  TORR.  &  GRAY,  Fl.  i :  33.     1838. 

D.  sarcophyllum  HOOK.  &  ARN.   Bot.  Beech.  317.      1841. 
D.  decorum   var.    nudicaule    HUTH,    Delph.    N.    Am.    9. 

1892. 
D.  -peltatum  HOOK.  ex.  Huth,  Bot.  Jahrb.  20:  449.     1895. 

Stem  i  to  T.y2  feet  high,  glabrous,  branched,  few-leaved: 
leaves  rather  succulent,  i  to  3  inches  across,  lobed  to  the  mid- 
dle or  farther  3  to  7  times,  the  secondary  lobes  rounded  and 
often  mucronate ;  petioles  3  to  5  inches  long,  dilated  at  the 
base :  flowers  panicled ;  sepals  bright  orange-red,  obtuse, 
scarcely  spreading,  shorter  than  the  stout  spur ;  petals  yellow, 
nearly  as  long  as  sepals ;  spurs  long  and  funnel-form  :  follicles 
3,  spreading  and  recurved,  soon  becoming  glabrous;  seeds 
thin-winged.  April  to  July.  Along  mountain  streams,  North- 
ern California.  Bot.  Mag.  5819.  Flor.  des  Serr.  19:  1949. 
Revue  Hort.  1893,  p.  259.  Marsh,  Hot  Springs  near  Santa 
Rosa,  Calif.,  a  pubescent  form  with  thicker  leaves.  Collected 
by  Coville,  May,  1884. 

4.  D.  cardinale  HOOK.  Bot.  Mag.  t.  4887.     1855. 
D.  coccineum  TORR.  Pac.  Ry.  Rep.  4:  62.      1857. 
D.jftammeum  KELLOGG,  in  Proc.  Calif.  Acad.  2  :  22.   1863. 

Stem  erect,  2  to  3^  feet  high,  partly  pubescent:  leaves 
smooth,  fleshy,  deeply  5-parted,  the  parts  cut  into  long,  linear 
lobes :  elongated,  many-flowered  raceme,  flowers  bright  red 
with  petal  limbs  yellow  :  follicles  glabrous,  usually  3  ;  seeds 
smooth.  July  to  Aug.  California.  Gartenflora,  208.  Flor. 
des  Serr.  u,  p.  63.  /.  7/05.  Garden  19:  273. 

5.  D.  viridescens  LEIBERG,  Proc.  Biol.  Soc.  Wash,  n  :  39. 
1897. 

Roots  fascicled  not  tuberous  :  plant  5  feet  high,  pubescent, 
especially  above  :  lower  stem-leaves  often  3-parted  and  again 
3~5-lobed  and  toothed ;  upper  leaves  dissected  into  narrow 
lobes ;  leaves  all  thin ;  pedicels  slender,  short,  appressed :  a 
narrow  bractlet  near  the  base  or  half  way  up,  and  a  pair  very 
near  the  flower  :  inflorescence  and  follicles  very  hairy  :  flowers 


Davis:   DELPHINIUMS  OF  NORTH  AMERICA.  437 

cream  to  greenish-yellow,  small;  spurs  nearly  horizontal, 
longer  than  the  sepals,  and  as  long  as  the  upper  pedicels. 
May  to  July.  Type  near  Peshaston  and  Wenatchee,  Okanogan 
Co.,  Wash.,  1500  feet  (f). 

6.  D.  Californicum  TORR.  &  GRAY,  Fl.  i :  31.     1838. 

D.  cxaltatum  HOOK.    &  ARN.    Bot.    Beech.    317.      1841. 

Not  Ait. 
D.  exaltatum  var.  Californicum  HUTH,  Delph.  N.  Am.  n. 

1892. 
D.  Californicum  var.  scapigerum  HUTH,  Bot.  Jahrb.  20 : 

451.     1895. 
?  D.  -viresccns  RYDB.  Bull.  Torr.  Club,  26:  385.     1899. 

(Fragment  only.) 

Stem  stout,  2  to  8  feet  high;  lower  leaves  very  large,  deeply 
cleft,  divisions  broad  wedge  shaped  ;  upper  with  narrower  divis- 
ions and  lanceolate  lobes  :  racemes  dense  :  flowers  sordid  whitish 
with  tinges  of  blue ;  sepals  and  spur  each  about  y±  inch  long  : 
follicles  much  like  those  of  D.  exaltatum.  Dry  places. 
Monterey  to  Mendocino  Co.,  Calif,  (f). 

Var.  laxiusculum  HUTH,  Bot.  Jahrb.  20:  451.     1895. 

Inflorescence  very  loose  and  open.  San  Francisco  region 
and  northern  Mexico. 

7.  D.  Przewalskii  HUTH,  Bot.  Jahrb.  20:  407.     1895. 
D.  Przewahkianum  HORT. 

Nearly  glabrous,  often  branched  at  base,  erect,  varying  much 
in  height :  leaves  3  to  5  times  deeply  parted  ;  parts  divided  into 
narrow  obtuse  lobes :  flowers  clear  yellow,  or  sometimes  tipped 
with  blue,  spur  equalling  the  sepals  :  follicles  3,  densely  hairy. 
July  to  Aug.  Asia. 

8.  D.  ZalilAiT.  &  HEMS.  Trans.  Linn.  Soc.  II,  3:  30.   1888. 
D.  hybridnm  var.  sulphureum  HORT. 

Stem  nearly  simple,  erect,  i  to  2  feet  high,  rather  glabrous 
or  becoming  so :  leaves  of  several  narrow,  linear  lobes,  dark 
green,  petioles  not  dilating  at  the  base :  flowers  large,  light 
yellow,  in  long  racemes:  follicles  3,  longitudinally  ribbed  and 
furrowed ;  seeds  with  transverse,  fibrous  plates.  June  to  July. 
Persia.  Bot.  Mag.  7049.  Garden  50:  1094;  54:  347. 
Gard.  Chron.  Ill,  20,  247. 


438  MINNESOTA    BOTANICAL    STUDIES. 

9.  D.  viride  WATS.  Proc.  Am.  Acad.  23:   268.     1888. 

Root  rather  thick,  branching  :  plant  glaucous,  about  2  feet 
high  ;  stems  glabrous  :  leaves  pubescent,  with  segments  acutely 
lobed,  upper  ones  more  deeply  divided  and  segments  narrower: 
racemes  open,  few-flowered  ;  pedicels  i  to  2  inches  long,  gla- 
brous or  somewhat  pubescent;  sepals  yellowish  green,  much 
shorter  than  the  stout  spur ;  petals  purple,  shorter  than  the  se- 
pals, lower  ones  entire  or  cleft,  villous  :  follicles  3,  not  spread- 
ing, very  finely  pubescent ;  seeds  large,  coats  dark,  wrinkled 
and  somewhat  winged  at  the  ends.  Gravelly  bluffs,  east  base 
of  Sierra  Madre,  Chihuahua,  Mex.  (f). 

10.  D.  bicolor  NUTT.  Journ.  Acad.  Phila.  7:   10.     1834. 
D.  Menziesii  GRAY,  Proc.  Acad.  Phila.  1863:  57.     Not 

DC. 

D.  Menziesii  var.    Utahense  WATS.  Bot.   King  Exp.  12. 
1871. 

Erect,  rather  stout,  ^  to  i  foot  high,  from  fascicled  roots  : 
leaves  small,  thick,  deeply  parted,  and  divisions  cleft  except 
perhaps  in  the  upper  leaves,  segments  linear :  obtuse  raceme 
rather  few-flowered  ;  the  lower  pedicels  ascending  i  to  2  inches  : 
spur  and  sepals  nearly  equal,  ^  inch  long  or  more,  blue ;  up- 
per petals  pale  yellow  or  white,  blue  veined  ;  lower  petals  blue  : 
follicles  glabrous  or  becoming  so.  May  to  Aug.  Dry  woods. 
Colorado,  west  and  north  to  Alaska. 

Var.  Montanense  RYDB.   Mem.   N.   Y.   Bot.   Gard.  i :   157. 
1900. 

Plant  glandular-pilose  ;  leaves  thicker  than  in  the  type.  Re- 
gion of  Helena  and  southward  into  Yellowstone  Park  (f). 

Var.  Nelsonii  n.  var.  D.  Nelsoni  GREENE,  Pitt.  3  :  92.     1896. 

Roots  sometimes  slightly  fascicled-tuberiform  :  lowest  leaves 
long-petioled  :  seeds  winged  as  in  the  type.  Southern  Wyom- 
ing to  middle  Colorado  (f). 

Var.  cognatum  n.  var. 

D.  cognatum  GREENE,  Pitt.  3:   14.     1896. 

Much  like  the  type  but  the  root  leaves  with  very  broad  seg- 
ments, plant  glabrescent,  or  hairy  on  the  flowers :  sepals  nar- 
rower than  the  type,  spurs  often  markedly  incurved  :  follicles 
3,  glabrous.  It  is  also  much  like  D.  Andersonii,  but  has  some 
stem  leaves,  and  the  flowers  are  different.  Western  Humboldt 
Mts.,  Nevada  (f). 


Davis:   DELPHINIUMS  OF  NORTH  AMERICA.  439 

Var.  glareosum  n.  var. 

D.  glareosum  GREENE,  Pitt.  3:   257.     1898. 

Rootstock  thick,  either  simple  or  branched :  plant  3  to  8 
inches  high,  with  i  to  3  stem  leaves:  follicles  3  to  5,  glabrous 
or  nearly  so.  Summit  of  Mt.  Steele,  Wash.  (f). 

n.  D.  decorum  FISCH.  &  MEY.  Ind.  Sem.   Hort.  Petrop.  3: 

33-     !837- 
D.  hesperium  HUTH,  Bot.  Jahrb.  20:  446.     1895. 

Stem  slender  and  weak,  ^  to  i  y2  feet  high,  smooth  or  nearly 
so  :  leaves  few,  bright  green,  upper  ones  small,  3~5~parted  into 
narrow  lobes,  lower  and  radical  ones  somewhat  reniform  in  out- 
line and  deeply  3-5-parted,  lobes  often  differing  widely :  flow- 
ers in  a  loose  raceme,  or  somewhat  panicled ;  sepals  blue,  *^ 
inch  long,  equalling  the  spurs  ;  upper  petals  at  least  tinged  with 
yellow  :  follicles  3,  thickish,  glabrous  ;  seeds  rugose,  not  winged. 
Spring.  Calif.  Bot.  Reg.  26:  64. 

Var.  gracilentum  n.  var. 

D.  gracilentum  GREENE,  Pitt.  3:   15.     1896. 

Differs  from  the  type  chiefly  in  the  radical  leaves,  which  are 
larger,  deeply  about  5-parted  or  lobed,  the  lobes  mostly  oval  or 
oblong,  obtuse  and  entire,  apiculate :  pedicels  often  filiform. 
Foothills  of  Sierra  Nevada  in  California  (f). 

Var.  patens  GRAY,  Bot.  Gaz.  12:  54.  1887. 
Z>.  -patens  BENTH.  PI.  Hartw.  296.  1848. 
D.  tricorne  var.  -patens  HUTH,  Delph.  N.  Am.  13.  1892. 

Stem  erect :  racemes  compact :  flowers  small,  sepals  a  third  to 
a  half  inch  long,  upper  petals  often  deeply  lined  with  blue ; 
seeds  somewhat  winged.  Siskiyou  Co.  to  southern  Califor- 
nia  (f). 

12.  D.  hesperium  GRAY,  Bot.  Gaz.  12:  54.     1887. 

D.  Jlfenziesuvar.  ochroleucum  TORR.  &  GRAY,  Fl.  1 :  31. 

1838. 

D.  azureum  TORR.  &  GRAY,  Fl.  I  :  660.     1840.     In  part. 
D.  azureum  &  D.  simplex  HOOK.  &  ARN.  Bot.  Beech. 

317.     1841. 
D.  simplex  WATS.  Bot.  Calif.  1 :   10.     1876. 

Roots  fascicled,  short,  some  of  them  fusiform,  2  feet  high ; 
stem  and  leaves  puberulent,  or  hairy  below  :  leaves  rather  small, 
much  dissected  into  narrow  parts  :  racemes  long,  many  flowered  ; 


440  MINNESOTA    BOTANICAL    STUDIES. 

flowers  violet- purple  varying  to  whitish,  sometimes  reddish 
purple ;  sepals  less  than  y2  inch  long,  about  equalled  by  the 
petals  and  by  the  spur ;  upper  petals  lined  and  bordered  with 
blue ;  pedicels  erect  in  fruit,  lowest  ones  about  i  inch  long, 
others  much  shorter:  follicles  3  to  5,  short-oblong,  puberulent, 
y%  inch  or  less  long ;  seeds  black  with  broad  light  wings  at  the 
angles.  West  Oregon  south  to  Monterey,  Calif,  (f). 

Var.  recurvatum  n.  var. 

D.  recurvatum  GREENE,  Pitt,  i:   285.     1889. 

Upper  petals  yellow,  not  bordered  nor  lined  with  blue.    Calif. 

(t)- 

13.  D.  Hanseni  GREENE,  Pitt.  3:  94.     1896. 

D.  hesperium  var.  Hanseni  GREENE,  Fl.  Fr.  304.     1892. 

D.  Hanseni  var.  arcuatum  GREENE,  Pitt.  3 :  94.  1896. 
Closely  allied  to  D.  hesperium,  but  very  slender :  racemes 
dense  but  lax  :  flowers  smaller  than  that  type  and  of  a  much 
lighter  blue ;  seeds  densely  scaly,  giving  a  white,  bur-like  ap- 
pearance. Amador  Co.,  Calif,  (f). 

14.  D.  variegatum  Torr.  &  Gray,  Fl.  i :  32.     1838. 

D.  grandiflorum   var.  variegatum  HOOK.  &  ARN.  BOT. 

Beech.  317.      1841. 
D.  decorum  BENTH.  PI.  Hartw.  295.      1848.    Not  Fisch. 

&  Mey. 

Root,  stem  and  leaves  like  D.  hes-perium:  flowers  larger,  only 
few  in  a  raceme  ;  sepals  much  surpassing  the  petals  :  follicles 
like  that  species  or  longer.  Monterey,  Calif.,  to  the  upper 
Sacramento  valley.  Common  along  streams,  etc.  Well  worth 
introduction  to  gardens  (f).  D.  Macounii  GREENE,  in  Herb. 
(Macoun  no.  18,078,  Geo.  Surv.  Canada)  is  a  low  weak  form 
or  variety  with  deeper,  less  fascicled  roots.  Rockies.  Lat. 

39°  40'.  " 

Var.  apiculatum  GREENE,  Fl.  Fr.  304.     1892. 
D.  apiculatum  GREENE,  Pitt,  i:   285.     1889. 
Leaf  segments  broader  :  flowers  more  numerous  (f). 
Var.  Blochmanae  n.  var. 

D.   ornatum  GREENE,  Fl.  Fr.  304.     1892.  ,  Not  Bouche. 
D.  Blochmance  GREENE,  Erythea,  i:   247.     1893. 
Leaf  segments  long   and  linear :   sepals   narrower  than  the 
type,  light  blue  or  white  ;    petals  with  crisp  margins.     Nipowa, 
Calif.     Specimens  at  Berkeley  (f). 


Davis :    DEEPHINIUMS  OF  NORTH  AMERICA.  441 

Var.  Emiliae  n.  var. 

D.  Emilice,  GREENE,  Erythea,  2:  120.      1894. 

Plants  often  3  feet  high  :  racemes  elongated  :  flowers  usually 
more  numerous  than  in  the  type.  Open  places  near  the  head 
of  Knight's  Valley,  Sonoma  Co.,  Calif,  (f). 

15.  D.  tricorne  MICHX.  Fl.  i :  314.     1803. 
D.flexuosum  RAF.  Ann.  Nat.  I  :  12.     1820. 

D.  aconitifolium  MUHLENB.  ex  Huth,  Bot.  Jahrb.  20 :  445. 

1895. 

Stem  succulent,  about  i  foot  high:  leaves  3-5 -parted  with 
3~5-cleft  linear  lobes ;  petioles  smooth,  hardly  dilating  at  the 
base  :  flowers  large,  blue,  rarely  whitish  ;  upper  petals  some- 
times yellow,  with  blue  veins ;  lower  ones  white-bearded ; 
sepals  nearly  equalling  the  spur :  follicles  3  or  4,  very  long, 
becoming  glabrous,  strongly  diverging ;  seeds  smooth.  May. 
Northern  States.  Lodd.  Bot.  Cab.  4:  306.  Very  beautiful 
and  much  used. 

16.  D   Andersonii  GRAY,  Bot.  Gaz.  12:  53.     1887. 

D.    decorum  var.   Nevadense   WATS.    Bot.   Calif,    i:    n. 

1876.     In  part. 

D.  Menziesii  WATS.  Bot.  King  Exp.     1871.     Not  DC. 
D.   tricorne  var.  Andersonii  HUTH,  Delph.  N.  Am.  13. 

1892. 

Stem  erect,  robust,  nearly  glabrous,  i^  feet  high  :  leaves 
rather  small,  thickish,  cuneate  divisions,  lobes  obtuse,  short: 
racemes  long,  dense :  flowers  blue ;  sepals  y2  inch  long,  shorter 
than  the  spur:  follicles  3  to  5,  about  yz  inch  long,  not  recurv- 
ing; seeds  winged.  Western  Nevada  to  mountains  of  Cali- 
fornia (f).  D.  Sonnet  GREENE,  Pitt.  3 :  264,  1897,  is  a 
slender,  weak  form,  from  California  (|). 

17.  D.  Parishii  GRAY,  Bot.  Gaz.  12:  53.     1887. 

Several  stemmed,  much  like  the  following,  but  with  racemes 
and  flowers  smaller :  sepals  oblong,  ^  to  ^  inch  long,  hardly 
surpassing  the  petals,  shorter  than  the  spur  ;  upper  petals  yel- 
lowish :  seed-coats  transversely  wrinkled ;  margin  broad,  wing- 
like.  Southeastern  California,  southward  into  Lower  Cali- 
fornia (f). 

18.  D.  Parryi  GRAY,  Bot.  Gaz.  12:  53.     1887. 

Much  like  the  last:  leaves  not  thick,  divisions  or  lobes  few 


442  MINNESOTA    BOTANICAL    STUDIES. 

and  linear :  sepals  oval,  over  y2  inch  long,  much  surpassing 
the  petals,  fully  as  long  as  the  spur :  follicles  as  in  the  last ; 
seeds  with  loose  coats,  folded  at  the  angles  forming  wing-like 
processes.  Southern  California  (f). 

19.  D.  Menziesii  DC.  Syst.  i  :  355.     1818. 

D.  paupercuium  GREENE,  Pitt,  i :  284.  1889. 
Plant  sparingly  pubescent:  stem  simple,  slender,  y2  to  i  l/> 
feet  high,  few-leaved  :  leaves  small,  3~5-parted,  the  divisions 
mainly  cleft  into  linear  or  lanceolate  lobes  ;  petioles  hardly  di- 
lating at  the  base  :  flowers  in  simple  conical  racemes ;  sepals 
blue,  somewhat  pubescent  outside,  nearly  equalling  the  spurs 
in  length;  upper  petals  yellowish:  follicles  3,  pubescent  or 
sometimes  glabrous ;  seeds  black-winged  on  the  outer  angles. 
April  to  June.  On  hills,  California  and  northward  to  Alaska. 
Bot.  Reg.  14 :  1192. 

20.  D.  pauciflorum   NUTT.  ex  Torr.  &  Gray,   FL    i:    33. 

1838. 

D.  Nuttallianum  PRITZ.  in  Walpers  Rep.  2  :  744.      1843. 
JD.   Menziesii    var.   -pauciflorum    HUTH,  Bot.  Jahrb.   20 : 

445-      1895. 

Stem  slender,  nearly  glabrous,  ^  to  i  foot  high ;  oblong  or 
fusiform  fasciculate-tuberous  roots :  leaves  small,  parted  into 
narrow  linear  lobes  ;  petioles  not  dilating  at  base  :  flowers  and 
fruit  similar  to  those  of  D.  Menziesii,  but  on  shorter  pedicels. 
May  to  June.  Colorado  to  Washington  and  California. 
Var.  Nevadense  GRAY,  Syn.  Fl.  i:  50.  1895. 

D.  decorum   var.  Nevadense   WATS.    Bot.  Calif,    i:    u. 

1876.     In  part. 

Leaves  much  dissected :  racemes  with  spreading  pedicels : 
flowers  often  pinkish  purple ;  sepals  longer  than  in  the  type 
but  shorter  than  the  spur:  follicles  much  like  the  type.  Sierra 
Nevadas,  above  Cisco,  and  in  Plumas  Co.,  Calif.,  into  Ne- 
vada (f). 

Var.  depauperatum  GRAY,  Bot.  Gaz.  12:    54.     1887. 

D.  tricorne  var.  depauperatum  HUTH,  Delph.  N.  Am.  13. 

1892. 

Stem  leaves  few,  lobes  ovate  to  lanceolate  :  racemes  fewer 
flowered  than  in  the  type  and  in  the  preceding  variety.  North- 
western Nevada  into  Oregon  (f). 


Da-vis :   DELPHINIUMS  OF  NORTH  AMERICA.  443 

21.  D.  Brunonianum  ROYLE,  111.  Bot.  Himal.  56.     1839. 
D.  moschatum  MUNRO  ex  Hook.  f.  &  Thorns.  Fl.  Ind.  53. 

1858. 

Stems  erect,  }4  to  i}4  feet  high  :  plant  somewhat  pubescent : 
upper  leaves  3-parted,  lower  ones  reniform,  5-parted,  segments 
deeply  cut,  musk  scented  :  flowers  large,  light  blue  with  pur- 
ple margins,  center  black;  spur  very  short;  sepals  i  inch  long, 
membranous  and  often  clinging  until  the  fruit  is  mature  :  folli- 
cles 3  or  4,  villose.  June  to  July.  China.  Revue  Belg.  1863  : 
34.  Bot.  Mag.  5461. 

22.  D.  Cashmirianum  ROYLE,  111.  Himal.  55.     1839. 

Plant  pubescent,  not  very  leafy :  stem  simple,  erect,  slen- 
der, 10  to  18  inches  high  :  root  leaves  orbicular,  2-3  inches  in 
diameter,  5~7-lobed,  coarsely  acutely  toothed  and  cut:  peti- 
oles 5-8  inches  long ;  stem  leaves  short-petioled,  3-5-lobed,  cut 
like  the  radical  ones,  all  rather  thick  and  bright  green :  inflo- 
rescence corymbose,  the  branches  rather  spreading  :  flowers  2 
inches  long,  deep  azure  blue ;  spur  broad,  obtuse,  inflated,  de- 
curved,  little  over  half  as  long  as  sepal ;  upper  petals  almost 
black,  2-lobed,  lateral  ones  greenish:  follicles  3  to  5,  hairy, 
July  to  September.  Himalayas.  Bot.  Mag.  6189.  Garten- 
flora,  1105.  Garden  18  :  261.  Rev.  Hort.  1893,  p.  259. 

Var.  Walkeri  HOOK.  Bot.  Mag.  t.  6830.     1885. 

Stem  very  short,  leafy,  many-flowered :  upper  leaves  less 
lobed  or  almost  entire,  small,  long-petioled  :  flowers  very  large, 
light  blue  with  yellow  petals.  Suited  to  rockwork. 

23.  D.  altissimum  WALLICH.  PI.  Asiat.  Rar.  2:  25.  t.  128. 
1831. 

Stem  tall  and  slender,  branched  ;  plant  shaggy-hairy  above  : 
leaves  palmately  5-parted,  the  divisions  3-lobed  and  toothed  ; 
bracts  long-lanceolate  :  flowers  blue  or  purple  in  long  branch- 
ing racemes  ;  spur  straight  or  slightly  incurved,  equalling  the 
sepals  ;  petals  2-lobed  :  3  erect  follicles  ;  seeds  not  winged  nor 
scaly.  August  to  September.  Himalayas. 

24.  D.  exaltatum  AITON,  Hort.  Kew.  i  ed.  2 :  244.     1789. 
D.  trydactylum  MICHX.  Fl.  i:  314.     1803. 
D.lilacinum    WIL.LD.    ex    Huth.    Bot.   Jahrb.    20:     455. 

1885. 
Stem  stout,  2  to  4  feet  high,  smoothish :   leaves  flat,  nearly 


444  MINNESOTA    BOTANICAL    STUDIES. 

glabrous,  deeply  cleft  into  3  to  7  wedge-shaped  lobes,  which 
are  often  trifid,  petioles  usually  not  dilated  at  the  base  :  flowers 
medium  in  size,  blue  with  upper  petals  yellow,  sepals  nearly 
equalling  the  spur  in  length ;  flowers  on  long,  crowded,  erect, 
pyramidal  racemes  :  follicles  3,  pubescent  or  smooth  ;  seed  coats 
irregularly  wrinkled.  June  to  August.  Borders  of  woods. 
Alabama  and  Carolina  to  Minnesota. 

25.  D.  Treleasei  B.  F.  BUSH,  n.  sp. 

Roots  fascicled,  rather  fleshy  :  stem  green,  slender  but  erect, 
usually  2  to  4  feet  high,  simple  or  branched,  glabrous  through- 
out, somewhat  glaucous  :  only  i  or  2  true  stem  leaves,  basal 
leaves  2  to  5,  2  to  5  inches  across,  about  5-parted,  and  lobed 
into  narrow  segments  with  calloused  tips  ;  petiole  of  lowest  stem 
leaf  dilating  :  inflorescence  open  and  somewhat  branching  ;  pedi- 
cels long  and  very  slender,  ascending  or  spreading,  pubescent  on 
upper  part ;  spurs  and  sepals  about  equal,  blue,  puberulent  with- 
out ;  petals  much  shorter  than  sepals  and  very  narrow ;  upper 
petals  blue  at  the  ends,  lower  ones  blue  with  dense  yellow  beard  ; 
spur  straight,  sometimes  2-lobed  :  sepals  narrowly  ovate  ;  bractlets 
very  small  and  slender,  usually  not  very  close  to  the  pubescent 
receptacle:  follicles  3,  not  divergent,  sparsely  hairy;  styles 
divergent,  %  the  length  of  follicles ;  stigmas  2-lobed ;  seed 
coats  dark  brown,  loose  and  much  wrinkled.  Collected  by  B. 
F.  Bush  (No.  73),  May  28,  1898,  Eagle  Rock,  Mo.  ;  (No.  81) 
June  10,  1899,  Forsyth,  Mo.  Common  in  barrens  (f). 

This  is  perhaps  most  nearly  related  to  D.  Carolinianum. 
Besides  the  difference  in  floral  characters  the  plant  is  glabrous, 
somewhat  glaucous :  roots  fleshy,  fascicled :  racemes  open : 
stem  leaves  about  2  :  follicles  always  3. 

26.  D.  Carolinianum  WALT.  Fl.  Carol.  155.     1788. 
D.  azureum  MICHX.  Fl.  I  :  314.      1803. 

D.  virescens  NUTT.  Gen.  2  :  14.     1818. 
D.  azureum  var.  laxiflorum  HUTH,  Bot.  Jahrb.  20  :   450. 
1895. 

Stem  i^  to  2^2  feet  high,  not  much  branched,  plant  some- 
what pubescent :  leaves  3~5-parted,  the  divisions  3~5-cleft 
into  usually  linear  lobes  :  spicate  racemes  slender,  usually  many- 
flowered  :  flowers  small,  azure-blue  ;  spurs  slender :  3  to  5  fol- 
licles, oblong,  erect ;  seeds  transversely  wrinkled  and  rough- 


Da-vis:   DELPHINIUMS  OF  NORTH  AMERICA.  445 

ened.     Florida  to  South  Carolina,  west  to  Missouri,  Arkansas, 
and  Mississippi.     Paxt.  Mag.  16 :  258. 

Yur.  album  HORT. 

A  garden  variety,  somewhat  taller :  leaves  larger  and  with 
broader  divisions  :  flowers  creamy- white.  There  is  a  double 
form  of  this  not  much  used  in  the  trade. 

Var.  vimineum  GRAY,  Bot.  Gaz.  12:  52.     1887. 

D.  vimineum  D.   DON,  in  Sweet's  Brit.  Fl.   Gard.  II,  4 : 

/.  374.     1838. 
D.  virescens  GRAY,  PI.  Lindh.  2  :  142.     1850. 

Very  slender  and  tall,  more  branched,  and  with  looser  inflo- 
rescence than  the  type :  seeds  larger,  transversely  winged  or 
deeply  and  thinly  wrinkled.  Gulf  region  of  Louisiana  and 
Texas.  Bot.  Mag.  3593.  Bot.  Reg.  13:  1999  (as  D.  azu- 
rcuni)  (f). 

27.  D.  Oreganum  HOWELL,  Fl.  N.  W.  Am.  i :  22.     1897. 

Tuber  flattish,  somewhat  branched :  plant  finely  pubescent, 
stem  often  slender,  i  to  2  feet  high,  sparingly  leafy  :  leaves  dis- 
sected into  acute  linear  lobes  :  racemes  rather  open :  flowers 
large,  blue ;  sepals  broadly  lanceolate,  shorter  than  the  slender 
spur,  and  longer  than  the  petals ;  upper  petals  yellow  or  white 
at  tip,  lower  ones  blue,  truncate,  bearded  :  follicles  3  to  4  lines 
long,  i  line  broad,  densely  tomentose,  not  spreading ;  seed  tri- 
angular with  rounded  and  rugose  back,  and  truncate  summit. 
Open  places.  Willamette  valley,  Oregon.  It  differs  from  D. 
CaroUnianum  chiefly  in  its  open  paniculate  inflorescence,  its 
very  small  follicles,  few  stem  leaves,  and  its  seed  characters  (f). 

28.  D.  Geyeri  GREENE,  Erythea,  2:  189.     Dec.,  1894. 

This  differs  from  D.  camportim  in  the  color  of  the  flowers, 
which  are  almost  wholly  blue,  and  in  having  the  upper  branch- 
lets  much  larger  than  in  that  species  :  seeds  somewhat  winged 
and  roughened.  High  plains,  western  Nebraska  and  Kansas, 
west  to  the  mountains  (f). 

Var.  Wootoni  n.  var. 

D.  Wootoni  RYDB.  Bull.  Torr.  Club,  26  :  587.     1899. 

This  southern  variety  is  intermediate  between  D.  cam^orum 
and  D.  Geyeri  in  the  size  of  its  upper  branchlets  :  sepals  blue  or 
bluish,  petals  white  or  nearly  so.  Arizona  and  New  Mexico  (f). 


446  MINNESOTA    BOTANICAL    STUDIES. 

Var.  geraniifolium  n.  var. 

D.  ger an i (folium  RYDB.  Bull.  Torr.  Club,  26  :  583.     1899. 

Differs  from  the  type  only  in  having  broader  leaf  segments, 
bractlets  variable  in  size,  and  pedicels  slightly  more  spreading. 
Charles  valley,  Arizona  (f). 

29.  D.  camporum  GREENE,  Erythea,  2:   183.     Nov.,  1894. 
D.  albescens  RYDB.  Bull.  Torr.  Club,  26:  583.      1899. 

Roots  fascicled,  fleshy-fibrous  :  stem  stout,  erect,  i  to  3  feet 
high,  pubescent  throughout,  especially  above  :  a  dense  cluster 
of  finely  dissected  root-leaves,  and  very  few  stem-leaves  :  ra- 
ceme long  and  simple,  often  dense ;  pedicels  short,  erect  or  ap- 
pressed  :  flowers  white  with  blue  spots  on  sepals,  and  sometimes 
tinged  with  blue  or  flesh  color ;  spurs  straight  or  curved,  longer 
than  the  sepals ;  upper  petals  often  tinged  with  yellow,  lower 
ones  2-lobed,  bearded  :  follicles  pubescent,  seeds  scaly  and  often 
winged  at  the  angles.  Widely  distributed.  Manitoba  to  Ar- 
kansas and  San  Antonio,  Texas,  west  to  the  Rockies  (f). 

Var.  Penardi  n.  var. 

D.  Penardi  HUTH,  in  Helios  10  :  27.     1893. 

Flowers  and  leaves  much  like  the  type  :  upper  petals  toothed  : 
seeds  large,  black,  slightly  scaly.  Flagstaff  Hill  and  Boulder, 
Colo.,j^  Huth.  No  type  of  this  is  known  in  America,  but 
seeds  of  it  have  been  sent  to  Columbia  University  by  M.  E. 
Autran,  of  the  Boissier  Herbarium.  A  specimen  from  Esmeralda 
Co.,  Neb.  (W.  H.  Shockley,  1881),  in  Gray  Herbarium  agrees 
in  characters  of  seeds  and  leaves,  but  not  in  color  of  flowers.  It 
is  an  intermediate  form  between  this  variety  and  D.  Geyeri  (f). 

Var.  macroseratilis  n.  var. 

D.  macroseratilis  RYDB.  Bull.  Torr.  Club,  26  :  585.    1899. 

Slender,  leaf-segments  fewer  than  in  the  t}'pe  :  flowers  much 
the  same.  Represents  the  southern  variation  of  the  camporum 
group.  Tom.  Greene  Co.,  Tex.  (f). 

30.  D.  scaposum  GREENE,  Bot.  Gaz.  6:  156.     1881. 

Root  a  cluster  of  thickened,  fleshy  fibres  :  stem  leafless  as  in 
D.  nudicaule;  radical  leaves  rather  fleshy,  pubescent,  3-parted, 
the  divisions  wedge-shaped,  3~5-cleft  or  toothed,  the  teeth  end- 
ing in  a  calloused  point :  racemes  many-flowered,  pedicels  as 
long  as  the  deep  azure  blue  flowers ;  spur  incurved :  follicles  3 
to  5  ;  seed  coat  somewhat  loose  and  wrinkled.  Southern  Utah 
and  Arizona  (f). 


Davis :   DELPHINIUMS  OF  NORTH  AMERICA.  447 

31.  D.  uliginosum  CURRAN,  Bull.  Calif.  Acad.  i :  151.  1885. 

Stem  leafless,  often  branching  :  radical  leaves  3-cleft,  lobes 
entire  or  i-3-toothed :  racemes  rather  few-flowered :  blue 
sepals  y&  inch  long,  equalling  the  straight  spur :  follicles  3  to 
5,  erect,  nearly  y2  inch  long;  seed  coats  minutely  wrinkled  and 
muriculate.  Lake  Co.,  Calif.,  in  swampy  ground  (f). 

32.  D.  trolliifolium  GRAY,  Proc.  Am.  Acad.  8:  375.     1872. 
D  exaltalum  var.  trolliifolium  HUTH,  Delph.  N.  Am.  u. 

1892. 

Stem  2  to  5  feet,  leafy,  often  reclining  :  leaves  thinnish,  large, 
often  reniform  at  base,  3~7-parted,  lobes  wedge-shape,  incised  : 
racemes  in  large  plants  i  to  2  feet  long  and  very  loose  :  flowers 
blue  with  upper  petals  white ;  spur  and  sepals  each  ^  inch 
long  :  follicles  glabrous ;  seeds  with  thin  wing  or  crown  at  the 
end.  April.  Moist  grounds,  Columbia  river. 

33.  D.  elatum  LINN.  Sp.  PI.  531.     1753. 

D.  intermedium  WILLD.  ex  Ait.  Hort.  Kew.  i  ed.  2 :  243. 

1789. 

D.  Clusianum  HOST.  Fl.  Aust.  2:  67.      1797. 
D.  alpinum  WALDST.  &   KIT.  PL    Rar.   Hung.    3:   273. 

1812. 

D.  -palmalifidum  DC.  Syst.  i  :  358.     1818  in  part. 
D.  ranunculi  folium  WALL.  Cat.  n.  4716.     1828. 
D.  pyramidale  ROYLE,  111.  Bot.  Himal.  56.     1839. 
D.  discolor  FISCH.  ex  Huth,  Bot.  Jahrb.  20:  399.     1895. 

Glabrous,  2  to  3^  feet  high:  leaves  somewhat  pubescent, 
5~7-parted,  parts  rather  narrow,  cut-lobed  ;  upper  leaves  3-5- 
parted ;  petioles  not  dilated  at  the  base :  raceme  much  like  D. 
exaltatum,  or  more  spike-like  :  flowers  blue  with  dark  violet 
petals  ;  sepals  ovate,  glabrous,  nearly  equalling  the  spurs  :  fol- 
licles 3,  seeds  transversely  wrinkled,  not  scaly.  June  to  Au- 
gust. Bot.  Reg.  23:  1963.  Gartenflora,  736  b  &  c  (vars.), 
Flor.  des  Serr.  12  :  1287  (var.  flore-pleno}.  Revue  Hort.  1859, 
p.  529  ;  1893,  p.  258. — A  polymorphous  and  complex  species  of 
Europe.  It  is  probable  that  all  or  nearly  all  the  plants  sold  here 
under  this  name  should  be  called  D.  exaltatum,  which  is  a 
closely  allied  species. 


448  MINNESOTA    BOTANICAL    STUDIES. 

34.  D.  grandiflorum  LINN.  Sp.  PI.  531.     1753. 

D.  sinense  FISCH.   ex  Link,    Enum.   Hort.   Berol.   2  :  80. 

1822. 
D.virgatum  JACQ^  f .    ex   Spreng.   Syst.    2:  617.      1825. 

Not  Poir. 

Stem  rather  slender,  2  to  3  feet  high :  leaves  rather  small, 
many  times  parted  into  nearly  distinct,  narrow,  linear  lobes  : 
flowers  large,  blue,  varying  to  white,  the  spur  and  lower  petals 
often  violet,  upper  petals  often  yellow ;  spurs  long  and  taper- 
pointed :  follicles  3,  pubescent;  seeds  triangular,  coats  wrin- 
kled, not  scaly.  July  to  August.  Siberia.  Bot.  Mag.  1686. 
Garden  46 :  /.  pp/  &  p.  484. 

There  are  several  garden  varieties :  var.  album,  HORT. 
Flowers  pure  white.  Var.  album-pleno  HORT.  Flowers  double 
and  pure  white.  Var.  jlore-ple:io  HORT.  Flowers  double, 
blue,  very  pretty. 

Var.  Chinensis  FISCHER  ex  DC.  Prod,  i :  53.     1824. 

Stems  very  slender,  not  much  branched  :  leaves  and  flowers 
like  the  type,  but  flowers  more  numerous.  China.  Lodd.  Bot. 
Cab.  i  :  71.  A  favorite  garden  plant. 

35.  D.  cheilanthum  FISCH.  ex  DC.  Syst.  i:  352.     1818. 
D.  magnificum  PAXT.  Mag.  Bot.  16 :   258.      1849. 
D.formosum  HORT.     Not  Boiss.  &  Huet. 

Stem  erect,  simple  or  branched,  2  to  3  feet  high  :  leaves  gla- 
brous or  slightly  pubescent,  5-parted,  the  lobes  pointed,  sub-tri- 
fid  and  somewhat  toothed :  flowers  dark  blue,  the  upper  petals 
sometimes  pale  yellow,  the  lower  ones  inflexed,  ovate,  entire ; 
spur  rather  long,  straight  or  somewhat  curved  :  3  follicles,  either 
glabrous  or  pubescent ;  seeds  three-cornered,  three-winged,  not 
scaly.  June  and  July.  Siberia.  Bot.  Reg.  6  :  473.  Garten- 
flora,  13:  253. 

36.  D.  scopulorum  GRAY,  PL  Wright.  2:  9.     1853. 
D.  exaltatum  HOOK.  Fl.  i:   25.     1829. 

D.  exaltatiim  var.  scopulorum  HUTH,  Delph.  N.  Am.  12. 

1892. 

Stem  2  to  5  feet  high,  glabrous  below :  leaves  5~7-parted, 
the  basal  ones  with  very  broad  segments,  which  are  round  and 
apiculate  at  apex ;  other  leaves  more  narrowly  cleft ;  petioles 
dilating  at  the  base  :  flowers  blue  or  purple,  rarely  white,  upper 


Da-vis:   DELPHINIUMS  OF  NORTH  AMERICA.  449 

petals  often  yellow  ;  spur  one-half  inch  long,  equalling  the  sepals  ; 
racemes  simple,  densely  many-flowered  :  follicles  3,  pubescent; 
seeds  black  with  loose   coats,  not  scary,  but  slightly  winged. 
Aug.  to  Sept.     Moist  ground,  west  of  Rockies. 
Var.  subalpinum  GRAY,  Bot.  Gaz.  12:  52.     1887. 
D.  occidcntale  WATS.  Bot.  Calif.  2  :  428.     1880. 
D.  elatum   var.   occidentale  WATS.  Bot.   King    Exp.    u. 

1871. 
D.  cxaltatum    var.  Barbeyi  HUTH,    Delph.  N.  Am.    n. 

1892. 

D.  Barbeyi  HUTH,  Bull.  Herb.  Boiss.  1 :  335.  t.  77.     1893. 
A   smaller   plant,    pubescent    above :    broader    divisions    of 
leaves  :  shorter  racemes  :  larger  and  deeper  colored  flowers  :  fol- 
licles glabrous  ;  seeds  much  like  the  type.    Wasatch  Mountains. 
Var.  stachydeum  GRAY,  Bot.  Gaz.  12:  52.     1887. 
Stem  erect,  3  to  6  feet  high  :  leaves  with  narrow  divisions  : 
plant    cinereous-pubescent   throughout ;    seeds  black  like    the 
type.     Oregon  to  New  Mexico  and  Arizona  (f). 

Var.  attenuatum  JONES,  Proc.  Cal.  Acad.  II,  5:  617.    1895. 
Stems  in  tufts,  3  to  4  feet  high  :  leaves  like  the  type  ;  .pubes- 
cence like  var.  subalpinum:  flowers  large,  deep  blue,  with  an 
odor  of  musk ;  sepals  long  and  narrow,  3  times  as  long  as  the 
petals,  and  longer  than  the  spur ;  upper  petals  white,  lower  ones 
bearded.     Utah.     Allied  to  D.  elatum  (f). 
Var.  diversifolium  n.  var. 

D.  diversifolium  GREENE,  Pitt.  3:  93.  1896. 
Stems  often  tufted,  rather  tall  and  slender :  lowest  leaves 
nearly  reniform  in  outline,  not  more  than  3-parted,  the  parts 
with  lobes  rounded  at  the  ends,  the  sinuses  very  narrow  ;  other 
leaves  like  the  type :  plant  somewhat  pubescent  in  upper  parts 
and  on  the  follicles.  Moist  meadows,  head  waters  of  the  Hum- 
boldt  river,  eastern  Nevada  (f). 

37.  D.  glaucum  WATS.  Bot.  Calif.  2:  427.  1880. 
D.  scopulorum  WATS.  Bot.  Calif.  I :  n.  1876. 
D.  scopulorum  var.  glaucum  GRAY,  Bot.  Gaz.  12:  52. 

1887. 
D.  exaltatum  var.  glaucum  HUTH,   Delph.  N.  Am.    n. 

1892. 

Much  like  D.  scopulorum:  plants  with  a  broader  type   of 
leaves,  often  glaucous,  glabrous,  or  the  pedicels  slightly  glan- 


450  MINNESOTA    BOTANICAL    STUDIES. 

dular-pubescent :  lower  petals  deeply  lobed  :  pistils  and  fruits 
glabrous  ;  seeds  black  with  light  wings.  Sierra  Nevada,  Cali- 
fornia and  San  Bernardino  mountains,  altitude  6,000  to  10,000 
feet,  north  to  Alaska  (f). 

38.  D.  glaucescens  RYDB.  Mem.  N.  Y.  Bot.  Gard.   i :   155. 
1900. 

Rootstock  thickened :  stem  somewhat  angled,  plant  finely 
pubescent  especially  above,  or  in  age  glabrate,  somewhat  glau- 
cous, i  to  2  feet  high :  leaves  divided  to  near  the  base  into  5  to 
8  cuneate  divisions,  these  generally  deeply  3-cleft :  raceme 
simple,  rather  short,  lower  bracts  linear,  longer  than  the  flowers, 
the  upper  ones  subulate ;  pedicels  and  flowers  densely  pilose, 
pedicels  spreading  :  flowers  dark  blue  or  variegated  with  white, 
somewhat  nodding ;  spur  straight,  equalling  the  sepals ;  upper 
petals  yellowish  white,  tipped  and  tinged  with  blue  :  ovaries 
densely  hairy  ;  fruit  not  seen.  Rocky  places,  Cedar  mountains, 
Montana,  and  Yellowstone  Park.  Differs  from  D.  glaucum  in 
its  shorter  and  more  pilose  inflorescence,  lower  and  more  tufted 
habit,  and  in  hairy  ovaries  (f). 

Var.  multicaule  RYDB.  Mem.  N.  Y.  Bot.  Gard.  i :  156. 
1900. 

More  bushy  than  the  type  and  less  pubescent :  leaf  segments 
longer  and  narrower :  flowers  smaller ;  spur  curved.  Rock 
slides,  Cedar  Mt.,  Montana  (f). 

39.  D.  Nuttallii  GRAY,  Bot.  Gaz.  12  :  54.     1887. 

D.    exaltatum  var.   Nuttattii  HUTH,    Delph.   N.    Am.   9. 

1892. 

D.  Columbianum  GREENE,  Erythea,  2:  193.      1894. 
D,  simplex  NUTT.  ex  Huth,  Bot.  Jahrb.  20:   472.      1895. 

Stem  erect,  simple,  nearly  glabrous,  leafy,  li  to  2^  feet  high  : 
leaves  thinnish,  3~5-parted,  parts  divided  into  many  linear-ob- 
long lobes  :  racemes  long,  many-flowered ;  sepals  deep  blue, 
ovate,  sparingly  pubescent,  shorter  than  the  spur ;  petals  blue 
or  upper  ones  yellow,  lower  ones  white-bearded:  follicles  3, 
pubescent,  rather  erect;  seeds  thin,  dark  with  yellow  wings. 
Summer.  Low  open  woods,  Columbia  River. 

Var.  leucophaeum  n.  var. 

D.  leucophceum  GREENE,  Erythea,  3:   118.     1895. 

A  slender  whitish  plant,  with  sepals  and  lower  petals  white, 
upper  petals  blue.  Oregon  (f). 


Da-vis  :   DELPHINIUMS  OF  NORTH  AMERICA.  451 

40.  D.  distichum  GEYER,  in  Hook.  London  Journ.  Bot.  6 : 
68.     1847. 

D.  simplex  var.  distichiflornm  HOOK.  1.  c.  67. 

D.  azurcum  TORR.  Bot.  Wilkes  Exped.  217.     1854. 

Not  so  tall  as  D.  simplex,  glabrous  or  inflorescence  pube- 
rulent :  leaves  rather  thicker  :  flowers  and  fruit  much  like  those 
of  D.  simplex;  upper  petals  whitish.  Low  prairies  of  eastern 
Oregon  and  Washington,  eastward  in  Montana  (f). 

Without  seeing  the  follicle  and  seeds  of  D.  Burkei  GREENE, 
Krythea,  2:  183,  1894,  it  is  best  not  to  consider  it  as  distinct 
from  D.  distichum.  The  type  specimen  is  at  Kew.  Thus  far 
no  preserved  specimens  showing  follicles  and  seeds  were  found. 
The  type  is  supposed  to  have  come  from  the  "  Snake  Country, 
probably  in  Idaho,"  but  collectors  in  that  region  have  been  un- 
able to  rediscover  the  plant. 

41.  D.  simplex  DOUGL.  ex  Hook.  Fl.  1 :  25.     1829. 

D.  azureum  var.  simplex  HUTH,  Delph.  N.  Am.  9.    1892. 

Stem  nearly  simple,  2  to  3  feet  high,  soft-pubescent  through- 
out :  leaves  many-parted  into  linear  divisions  and  lobes : 
racemes  dense,  little  branched :  flowers  pale  blue  with  upper 
petals  yellow,  lower  petals  white-bearded ;  sepals  equalling  the 
spur:  follicles  3,  pubescent;  seeds  dark  with  margins  white- 
winged.  June.  Mountains  of  Idaho  and  Oregon. 

42.  D.  formosum  Boiss.  &  HUET,  Diagn.  Sec.   II,  5:    13. 

1856. 
D.  speciosum  Boiss.  &  HUET  ex  Huth,  Bot.  Jahrb.    20  : 

410.     1885.     Not.  M.  Bieb. 

Stem  strong,  2  to  3  feet  high,  hairy  below,  rather  glabrous 
above  :  lower  leaves  5~7-parted,  long-petioled,  upper  ones  3-5- 
parted,  short-petioled  or  sessile,  all  alternate :  racemes  many- 
flowered  :  flowers  blue  with  indigo  margins ;  spur  long,  violet 
colored,  bifid  at  the  tips :  follicles  3,  pubescent;  seeds  scaly. 
June  to  July.  Asia  Minor,  perhaps  ;  but  its  origin  is  disputed. 
Flor.  des  Serr.  12  :  1185. 

43.  D.  Maackianum  REGEL,  in  Mem.  Acad.  Petersb.  VII, 
4:  9.     1861. 

Erect,  3  feet  high,  pubescent  or  glabrous,  branched  above : 
leaves  pubescent  on  both  sides,  base  often  truncate  or  reniform, 
3-5-parted,  the  parts  serrate;  the  bases  of  petioles  dilated: 


452  MINNESOTA    BOTANICAL    STUDIES. 

flowers  in  loose  panicles ;  peduncles  yellow-hairy,  with  the 
bracts  often  inserted  above  the  base ;  sepals  blue,  y2  as  long  as 
the  spurs  ;  petals  dark  violet :  follicles  often  glabrous,  ^  inch 
long ;  seeds  small,  distinctly  scaly.  July.  Siberia.  Garten- 
flora,  344. 

44.  D.  hybridum  STEPH.  ex  Willd.  Sp.  PI.  2:   1229.     1799. 
D.  davuricum  GEORGI,  Beschr.  Russ.  Reich.  Ill,  4  :  1052. 

1800. 
D.fissum  WALDST.  &  KIT.  PI.  Rar.   Hung.  I  :  83.  /.  81, 

1802. 

D.  hirsutum  PERS.  Syn.  2:  82.      1807. 
D.  tauricum   PALLAS  ex  Bieb.    Fl.    Taur.   Cauc.   2:   13. 

1808. 

Stem  3  to  4  feet  high,  pubescent  above  :  root  somewhat  bulb- 
ous :  leaves  5-  to  many-parted,  lobes  linear,  petioles  dilated  and 
sheathing  at  the  base  :  racemes  dense  :  flowers  blue,  lower  limbs 
white-bearded ;  straight  spur,  longer  than  the  sepals  :  follicles 
3,  hairy;  ovate  seeds  with  transverse  scales.  June  to  August. 
Mountains  of  Asia.  Revue  Hort.  1893,  p.  258. — There  are 
many  double  and  semi-double  varieties  of  this  type.  Var.Jtore- 
•pleno  HORT.,  has  large  double  flowers  colored  as  in  the  type. 
Var.  Barloivi  PAXT.,  has  very  large  semi-double  flowers,  deep 
blue  with  brownish  center ;  a  supposed  hybrid  with  D.  grandi- 
jlorum.  Bot.  Reg.  1944. 

45.  D.  Madrense  WATS.  Proc.  Am.  Acad.  25:  141.     1890. 

Resembles  D '.  -pauciflorum ;  slender,  2  feet  high  or  less,  from 
a  thickened  rootstock,  pubescent  with  reflexed  hairs  below, 
glandular-hispid  above  :  leaves  3-parted,  the  lobes  5-7 -cleft  into 
linear-oblong  segments,  lowest  ones  less  cleft :  flowers  few, 
small,  pale  blue,  in  a  slender  raceme;  spur  narrow,  straight; 
lateral  petals  long-villous :  carpels  short,  glandular-hispid. 
May.  In  mountains  near  Monterey,  Mexico.  Collected  by 
Pringle  (n.  3014)  (f). 

46.  D.  bicornutum  HEMSL.  Diagn.  PL  Nov.  2:   17.     1879. 

Stem  nearly  simple,  stout,  2  to  4  feet  high,  glabrous  or 
puberulent  on  inflorescence  and  under  the  leaves  :  leaves  long- 
petioled,  5-parted,  3~5-lobed ;  bracts  linear ;  pedicels  bibrac- 
teolate ;  bractlets  cuneate  :  flowers  blue,  spurs  nearly  straight, 


Davis :   DELPHINIUMS  OF  NORTH  AMERICA.  453 

sepals  oblong  ;  upper  petals  narrow,  obtuse  or  retuse ;  lower 
petals  bifid :  carpels  3,  puberulent  at  first.  Oaxaca  (f). 

Var.  Hemsleyi  HUTH,  Bot.  Jahrb.  20:  453.     1895. 

Spurs  distinctly  2-lobed  at  the  end  (f). 

47.  D.  Ehrenbergi   HUTH,  in  Bull.    Herb.  Boiss.    i:  336.  t. 

//,/.   2.        1893. 

Stem  simple,  succulent,  very  leafy,  i  to  2  feet  high ;  petioles 
long  with  dilating  bases  :  leaves  3~5-parted,  cut  into  many  ob- 
long or  linear  lobes  :  racemes  few-flowered,  pedicels  erect,  lower 
ones  i  to  2  inches  long  :  flowers  blue ;  spurs  straight,  equalling 
the  sepals,  which  are  oval,  pubescent;  upper  petals  yellow  with 
blue  tips,  lower  ones  2-lobed,  sparsely  bearded,  appendiculate 
at  base  :  follicles  3,  erect,  pubescent.  Near  El  Cerro,  Mex.  (f.). 

48.  D.  pedatisectum  HEMSL.  Diagn.  PI.  Nov.  2:  18.    1879. 

Stem  branching,  branches  smooth  :  leaves  3~7-parted,  parts 
scarcely  lobed,  puberulent :  flowers  blue,  on  long,  slender, 
puberulent  pedicels ;  bracts  and  bractlets  linear-subulate ; 
sepals  oblique-oblong ;  upper  petals  deeply  2-lobed,  much 
shorter  than  the  spur,  lower  petals  2-lobed,  bearded,  appendi- 
culate at  base;  stigmas  glabrous:  follicles  3,  tomentose  when 
young  ;  styles  long.  Mexico.  Specimen  at  Kew  (t). 

49.  D.  latisepalum  HEMSL.  Diagn.  PI.  Nov.  2:   17.     1879. 

Plant  pubescent  or  villose  :  stem  nearly  simple :  leaves  5- 
parted ;  the  parts  of  the  basal  leaves  again  3~5-lobed,  parts  of 
stem  leaves  nearly  linear :  flowers  few  on  slender  pedicels ; 
spurs  slightly  curved,  nearly  equalling  the  sepals ;  sepals  vil- 
lose ;  upper  petals  narrow,  slightly  2-lobed  at  the  points  ;  lower 
petals  not  appendiculate,  deeply  2-lobed,  much  bearded  on  both 
sides;  stamens  glabrous:  follicles  3,  clothed  with  white 
pubescence  when  young.  9,000  feet.  Mt.  Tanga,  Oaxaca, 
Mex.  Specimen  at  Kew  (f). 

50.  D.  tenuisectum  GREENE,  Erythea,  2 :   184.     1894. 
Root  thick,  woody,  deep  :  stem  2  to  3  feet  high,  simple  or 

little  branched,  not  very  stout,  sulcate  above :  plant  finely 
pubescent  throughout,  leafy  :  leaves  very  finely  dissected  into 
linear  segments,  the  lower  stem  leaves  on  rather  long  petioles 
dilated  at  base  :  racemes  about  8  inches  long,  loosely  flowered, 
pedicels  very  slender,  nearly  erect,  upper  ones  not  longer  than 


454  MINNESOTA    BOTANICAL    STUDIES. 

the  spurs  ;  bractlets  of  lower  pedicels  lobed  ;  bractlets  of  upper 
pedicels  slender  and  near  the  flowers ;  sepals  about  equal  to 
spur  in  length,  blue  within,  tinged  with  yellow  outside  ;  upper 
petals  yellow  with  blue  tips  ;  lower  ones  either  blue  or  yel- 
lowish, 2-lobed  with  a  few  long,  white  hairs  on  inside  of  lobes : 
follicles  3,  large,  slightly  spreading;  seeds  nearly  black,  coats 
roughened,  forming  slight  wings  at  the  angles.  Cool  banks  of 
ravines  in  plains  at  base  of  the  Sierra  Madre,  Chihuahua,  Mex. 
Collected  first  by  C.  G.  Pringle  (n.  1184),  Sept.  27,  1887. 
Differs  essentially  from  D.  scopulorum  in  its  finely  dissected 
leaves  (f). 

51.  D.  leptophyllum  HEMSL.  Diagn.  PI.  Nov.  2  :  18.     1879. 

Stem  3  to  4  feet  high,  glabrous,  somewhat  branched  :  leaves 
glabrous,  deeply  5-parted,  and  cut  into  oblong  to  linear  lobes ; 
bracts  entire,  linear  :  inflorescence  open,  few-flowered  ;  pedicels 
i  to  2  inches  long  ;  bractlets  remote  from  flower  :  flowers  large, 
blue ;  spur  large ;  sepals  puberulent,  ovate,  obtuse,  2/$  inch 
long ;  petals  dull  yellow ;  upper  ones  slightly  2-lobed,  nearly 
glabrous ;  lower  ones  deeply  2-lobed,  slightly  bearded  :  folli- 
cles 3,  densely  villose  when  young,  half  inch  long  when  mature, 
not  spreading  :  seed  slightly  winged,  and  transversely  wrinkled. 
October.  San  Luis  Potosi  and  Montes  San  Miguelito,  Guana- 
juata,  southern  Mexico  (f). 

52.  D.  Wislizeni  ENGELM.   in  Wisliz.   Tour  N.   Mex.    106. 
1848. 

Stem  simple,  2  to  2)4  feet  high,  slender,  glabrous,  glaucous; 
petioles  elongated,  lower  ones  dilated  at  base  :  leaves  cut  into 
linear  segments;  pedicels  long:  flowers  few,  spur  2^  inches 
long,  blue,  slightly  pubescent  outside ;  the  outer  sepal  acute, 
others  obtuse  :  follicles  glabrous  even  when  young.  Wislizeni 
region,  Mexico.  Later  found  near  Cosihuiriachi,  8,000  feet. 
In  flower  in  Sept.  (f). 

EXCLUDED. 

D.  urceolatum]hco^  Coll.  I:  153,  1786,  is  figured  in  Bot. 
Mag.  n.  I791*  but  no  nativity  is  given.  From  character  it  may 
be  allied  to  D.  exaltatum,  but  it  is  probably  not  American. 


Davis:   DELPHINIUMS  OF  NORTH  AMERICA. 


455 


INDEX   TO    SPECIFIC  AND   VARIETAL    NAMES    OF 
DELPHINIUM. 


mconitifolium,  15. 

addendum,  \ . 

Ajacis,  i. 

albesccns,  29. 

alpinum,  33. 

altissimum,  23. 

Andersonii,  16. 

appiculatum,  14. 

azureum  Hook.  &  Arn.,   12. 

azureum  Michx.,  26. 

azureum  Torr.,  40. 

azureum  T.  &  G.,  12. 

azureum  var.  laxiflorum,  26. 

azureum  var.  simplex,  41. 

Barbeyi,  36. 

bi color,  10. 

bicolorvur.  cognatum,  10. 

bicolor  var.  glareosum,  10. 

bicolor  var.  Montanense,  10. 

bicolor  var.  Nelsonii,  10. 

bicornutum,  46. 

bicornutum  var.  Hemsleyi,  46. 

Blochmance,  14. 

Brunonianum,  21. 

Burkeyi,  40. 

Californicum,  6. 

Californicum  var.   laxiusculum, 
6. 

Californicum  var.    scapigerum, 

6. 

camporum,  29. 
camporum    var.     macroscratilis, 

29. 
camporum  var.  Penardi,  29. 

cardinale,  4. 

Carolinianum,  26. 

Carolinianum  var.  album,  26. 

Carolinianum    var.     Tjimineum, 
26. 

Cashmirianum,  22. 


Cashmz'rianumvar.  Walkeri,  22. 
CERATOSANTHUS  Aj'acis,  i. 
CERATOSANTHUS  consolida,  2. 
cheilanthum,  35. 
clusianum,  33. 
coccineum,  4. 
cognatum,  10. 
Columbianum,  39. 
consolida  Sibth.   &  Sm.,  i. 
consolida  Linn.,  2. 
davuricum,  44. 
decorum  Benth.,  14. 
decorum  Fisch.  &  Mey.,  n. 
decorum  var.  gracilentum,  1 1 . 
decorum  var.  Nevadense,  16  and 

20. 

decorum  var.  nudicaule,  3. 
decorum  var.  patens,  1 1 . 
discolor,  33. 
distichum,  40. 
diversifolium,  36. 
Ehrenbergi,  47. 
elatum,  33. 

elatum  var.   occidejttale,  36. 
Emilice,  14. 
exaltatum  Ait.,  24. 
exalt atum  Hook.  &  Arn.,  6. 
exaltatum  Hook.,  36. 
exaltatum  var.  Barbeyi,  36. 
exaltatum  var.   Californicum,  6. 
exaltatum  var.  glaucum,  37. 
exaltatum  var.  Nuttallii,  39. 
exaltatum  var.  scopulorum,  36. 
exaltatum  var.  trolliifolium,  32. 
Jissum,  44. 
jlammeum,  4. 
Jlexuosum,  15- 

formosum  Boiss.  &  Huet,  42. 
formosum  Hort.,  35. 
geraniifolium,  28. 


456 


MINNESOTA    BOTANICAL    STUDIES. 


Geyeri,  28. 

Geyeri  var.  geraniifolittm,  28. 
Geyeri  var.    Wootoni,  28. 
glareosum,  10. 
glaticescens,  38. 
glaucescens  var.  multicaule,  38. 
glaucum,  37. 
gracilentum,  \  i . 
grandiflorum,  34. 
grandiflorum  var.  Chinensis,  34. 
grandiflorum   var.    variegatum, 

14. 

Hanseni,  13. 

Hanseni\n.r.  arcuatum,  13. 
kesperium  Huth,  1 1 . 
hesperium  Gray,  12. 
hesperium  var.  Hanseni,  13. 
hesperium  var.  recurvatum,  12. 
hirsutum,  44. 
hybridum,  44. 
hybridum  var.  Barlo'wi,  44. 
hybridum  var.  sulphureum,  8. 
intermedium,  33. 
latisepalum,  49. 
leptophyllum,  51. 
leucophceum,  39. 
lilacinum,  24. 
Maackianum,  43. 
Macounii,  14. 
macroseratilis,  29. 
Madrense,  45. 
magnificum,  35. 
Menziesii  DC.,  19. 
Menziesii  Wats . ,  1 6 . 
Menziesii  Gray,  10. 
Menziesii  var.  ochroleucum,  12. 
Menziesii  var.  pauciftorum,  20. 
Meuziesii  var.   Utahense,  10. 
monophyllum,  2. 
moschatum,  21. 
Nelsoni,  10. 
midicaulc,  3. 
Nuttallii,  39. 


N2ittallii  var.  leucophcetim^,  39. 

Nuttallianum,  20. 

occidentale,  36. 

Oreganum,  27. 

ornatum  Bouche,  i . 

ornatum  Greene,  14. 

palmatifidum,  33. 

Parishii,  17. 

Parryi,  18. 

patens*  1 1 . 

pauciflorum,  20. 

paucijlorum  var.  depauperatum, 

20. 

paucijlorum  var.  Nevadense,  20. 
pauperculum,  19. 
pedatisectum,  48. 
peltatum,  3. 
Penardi,  29. 
Przewalskianum,  7. 
Przeijoalskii,  7. 
pubescens,  I. 
pyramidale,  33. 
r a « tinctdifoliu m,  33. 
recurvattim,  12. 
5«  rcophyllum , '  3 . 
scaposum,  30. 
scopulorum  Gray,  36. 
scopidorum  Wats.,  37. 
scopulorum  var.  attenuatum,  36 
scopulorum    var.     divers  if olium, 

36. 

scopulorum  var.  glaucum,  37. 
scopulorum  var.  stachydeum,  36. 
scop2ilorum  var.  subalpinum,  36. 
segetum,  2. 
simplex  Dougl.,  41. 
simplex  Hook.  &  Arn.,  12. 
simplex  Nutt.,  39. 
simplex  Wats.,  12. 
simplex  var.  distichijlorum,  40. 
sinense,  34. 
Sonnei,  16. 
speciostim,  42. 


DELPHINIUMS    OK    NORTH    AMERICA. 


457 


tauricum,  44. 
tenuisectum,  50. 

Trclcasei,  25. 

tricorne,  15. 

tricorne  var.  Andersonii,  16. 

tricorne  var.   depauperatum,  20. 

tricorne  var.  patens,  II. 

trolliifolium,  32. 

trydactylum,  24. 

tiliginosum,  3 1 . 

•uariegatum,  14. 

•variegatum  var.  apiculatum,  14. 

varicgatiim  var.  Blochmance,  14. 


variegatum  var.  E/nilice,  14. 
versicolor,  2. 
vimineum,  26. 
virescens  Gray,  26. 
virescens  Nutt.,  26. 
virescens  Rydb.,  6. 
virgatum,  34. 
viride,  9. 
viridescens,  5. 
Wootoni,  28. 
Wislizeni,  52 • 
z7,  8. 


XXVI.    NATIVE  AND  CULTIVATED  RANUNCULI  OF 
NORTH  AMERICA  AND  SEGREGATED  GENERA. 


K.  C.  DAVIS. 


Few  groups  of  plants  are  perhaps  less  understood  than  this 
one.  The  range  of  variation  in  characters  is  rather  limited  and 
yet  the  number  of  species  is  very  great.  Six  genera  have  taken 
from  Ranunculus  about  thirty-six  species  and  yet  that  genus  re- 
tains more  species  than  any  other  of  the  order  Ranunculaceae. 
About  350  names  have  already  been  given  to  American  plants 
of  this  group. 

KEY  TO  GENERA. 

A.  Akencs  transversely  wrinkled ;    roots    fibrous ;    aquatic    or   ditch 

herbs;  flowers  white Batrachium. 

AA.   Akenes  not  transversely  wrinkled. 

B.   Developed  carpels  not  longitudinally  ribbed  or  striated. 

C.  Roots  not  a  cluster  of  thickened  tubers,  or  several  times  longer 

than  thick. 
D.  Flowers  mostly  yellow  or  white;  akenes  compressed,  never 

lanceolate,  srgooth,  papillose  or  spiny Ranunculus. 

DD.  Flowers  white;  akenes  lanceolate,  utricular;  style  hooked. 

Kumlienia. 
CC.  Roots  a  cluster  of  thickened  tubers;  leaves  crenate,  cordate; 

cotyledon  only  i Ficaria. 

BB.   Developed  carpels  longitudinally  ribbed  or  striated. 

C.   Leaves  pinnately  compound   or    lobed ;   akenes  terete,    style 

persistent,  slender,  recurved Cyrtorhyncha. 

CC.   Leaves  not  as  above ;  akenes  compressed. 

D.  Akenes  with  beaks  somewhat  reflexed ;  leaves  rounded  and 

lobed Arcteranthis. 

DD.  Akenes  minutely  sharp-pointed;   leaves    crenate-dentate, 

oval-cordate  to  renif orm Oxygraphis. 

459 


460  MINNESOTA    BOTANICAL    STUDIES. 

BATRACHIUM  S.  F.  GRAY,  Nat.  Arr.  Brit.  PI.  2:  720. 

1821. 

(Name  from  the  Greek,  in  allusion  to  the  aquatic  habitat  of 
the  plants.) 

Aquatic,  or  semi-aquatic  perennial  herbs  ;  leaves  dissected  or 
lobed,  submerged  ones  usually  with  filiform  segments  ;  petioles 
with  stipular-dilated  membranous  bases  :  flowers  solitary,  op- 
posite the  leaves,  rather  small,  white ;  sepals  usually  5  ;  petals 
usually  5,  base  often  yellowish  ;  claw  with  a  naked  nectar  pit; 
stamens  several  or  many :  ovules  oblique,  compressed,  not 
margined,  nearly  beakless,  transversely  rugose.  About  20 
species,  mostly  of  north  temperate  regions  of  the  world.  The 
following  are  all  that  are  found  in  North  America.  Section 
BATRACHIUM  DC.  Syst.  I  :  233,  under  Ranunculus. 

KEY  TO   SPECIES. 

A.  Aquatic  leaves  with  filiform  segments  present ;  receptacle  hairy. 

B.  Leaves  all  sessile  or  nearly  so i.   divaricatum. 

BB.  Leaves,  except  the  upper  ones  long-petioled. 

C.  Emersed  leaves  always  present,  with  segments  broader  than 

linear 2 .  aquatile. 

CC.  Emersed  leaves  if  present  only  fleshy  or  nearly  linear. 

3.   trichophyllum. 

AA.  Aquatic  leaves  none,  or  few,  and  with  few  divisions ;  receptacle 
glabrous. 

B.   Styles  minute,  shorter  than  the  ovaries 4.   hederaceum. 

BB.  Styles  long  and  filiform 5.   Lobbii. 

i.  B.  divaricatum  WIMM.  Fl.  Schles.  10.     1841. 
Ranunculus  aquatilis  /9.  LINN.  Sp.  PL  556.     1753. 
R.  divaricatus  SCHRANK,  Baier.  Fl.  2:   104.     1789. 
R.  circinalus  SIBTH.  Fl.  Oxon.  175.     1794- 
7?.  aquatilus  var.  stagnatalis  DC.  Prod.   I  :   27.      1824. 
Batrachium  circinatum  SPACH.  Hist.  Veg.  7*.  201.      1839. 
R.  stagnatalis  WALLR.  Sched.  Crit.   285.      1848. 
R.  aquatilis  var.  divaricatus  GRAY,  Man.  2  ed.  7.    1856. 
R.  longirostris  GODRON,  Ess.  32.  f.  9.     1862. 
R.  aquatilis    var.     longirostris     LAWSON,     Rev.    Canad. 
Ranunc.  43.      1870. 

Leaves  sessile  to  the  dilated  stipule-like  base,  dissected  into 
rigid  lobes  spreading  at  right  angles  to  the  stem,  not  collapsing 
when  taken  from  the  water ;  no  floating  nor  emersed  leaves : 


Davis  :   RANUNCULI  OF  NORTH  AMERICA.  461 

petals  several-nerved,  deciduous  :  styles  subulate,  as  long  as  the 
ovaries,  stigma  surface  along  the  inner  side  :  receptacle  hairy. 
July.  Chihuahua,  Mex.,  Texas  to  British  Columbia,  eastward 
and  northward  to  Hudson  Bay.  Also  in  Europe. 

2.  B.  aquatile  WIMM.  Fl.  Schles.  8.     1841. 
Ranunculus  aquatilis  LINN.  Sp.  PI.  556.      1753. 

R.  aquatilis  var.  heterophyllus  DC.  Prod.  I  :   26.      1824. 
R.  aquatilis  var.  hispidulus  DREW,  Bull.  Torr.  Club,  16 : 

150.      1889. 
R.    Grayanus   FREYN,   Deutsche    Bot.   Monats.    8:   179. 

1891. 

Floating  leaves  round-reniform,  3-5-lobed  or  parted,  and 
the  divisions  2— 3-cleft ;  submersed  ones  with  filiform  segments, 
widely  spreading,  rather  firm,  but  collapsing  when  taken  from 
water ;  all  the  leaves  often  slightly  hispid  below  :  styles  sub- 
ulate, shorter  than  the  ovaries,  introrsely  stigmatose  :  recep- 
tacle hairy  among  the  carpels.  Ponds  and  quiet  shallow 
streams.  California  to  Alaska,  Europe  and  Asia. 

3.  B.  trichophyllum  BOSSCH.  Prod.  Fl.  Bot.  5.     1850. 
Ranunculus  trichophyllus  CHAIX.  in  Vill.  Hist.  PL  Dauph. 

i:  335.     1786. 
R.  jlaccidus  PERS.   in   Usteri.  Ann.  Bot.  5 :  pt.   14 :  39. 


R.  pantothrix  BROT.  ex  DC.  Syst.  i :  235.     1818. 

7?.  aquatilis  var.  ccespitosus  DC.  Prod.  I  :   26.     1824. 

R.  aquatilis  var.  brachypus  HOOK.   &   ARN.    Bot.   Beech. 

316.     1841. 

R.  confer-voides  FRIES,  Sum.  Veg.  Scand.  1 :   139.     1846. 
R.  aquatilis  var.  submersus    GORDON,  in   Gren.   &   Godr. 

Fl.  Fr.  i  :   23.     1848. 

R.  aquatilis  var.  trichophyllus  GRAY,  Man.  5  ed.  40.    1867. 
R.  Porteri  BRITTON,  Bull.  Torr.  Club,  17:  310.     1890. 
R.  aquatilis  var.  confervotdesGRAY,  Syn.  Fl.  1:21.    1895. 
R.  aquatilis  v&r.Jlaccidus  GRAY,  1.  c.  21. 

This  species  is  polymorphous,  including  those  with  filiform 
segments  to  all  the  leaves  or  with  some  of  the  leaves  rather 
fleshy,  or  some  narrowly  linear,  and  the  submersed  leaves  are 
mostly  flaccid,  but  ^ay  be  rigid  when  taken  from  the  water. 
The  plants  have  adapted  themselves  to  either  aquatic  or  to 
muddy  habitats.  Widely  distributed  ;  America,  Europe,  Asia. 


462  MINNESOTA    BOTANICAL    STUDIES. 

4.  B.  hederaceum  S.  F.  Gray,  Nat.  Arr.  Brit.  PI.   2:  721. 

1821. 
Ranunculus  hederaccus  LINN.  Sp.  PL  556.     1753. 

Semi-aquatic,  rooting  freely  from  the  nodes,  in  the  mud : 
leaves  seldom  submersed,  but  floating  or  resting  on  the  mud ; 
all  reniform  or  nearly  so,  angulate-lobed,  never  finely  dissected  ; 
peduncles  as  short  as  the  petioles :  petals  deciduous :  styles 
shorter  than  ovaries,  introrsely  stigmatose ;  receptacle  glabrous. 
Naturalized  from  Europe  at  Norfolk,  Va.,  and  on  Newfound- 
land. 

5.  B.  Lobbii  HOWELL,  Fl.  N.  W.  Am.  i:    13.     1897. 
Ranunculus    hederaceus  var.   TORR.   Pac.    Ry.   Rep.   4 : 

62.     1853. 
7?.  hederaceus  var.  Lobbii  LAWSON,  Rev.  Canad.  Ranunc. 

44.     1870. 
R.  hydrocharis  subsp.   Lobbii  HIERN.  Seem.  Journ.  Bot. 

9:  66.  /.  114.     1871. 

R.  aquatilis  var.  Lobbii  WATS.  Bibl.  Index  17.     1878. 
R.  Lobbii  GRAY,  Proc.  Am.  Acad.  21  :  364.      1886. 

Leaves  commonly  all  floating,  small,  truncate  or  cordate  at 
base,  divergently  3-parted :  petals  persistent ;  stamens  5  to  10  ; 
styles  long  and  filiform;  stigma  terminal.  In  mud  or  water  of 
pools,  etc.  California  and  Oregon. 

RANUNCULUS  LINN.  Sp.  PL  548.     1753. 

The  name  is  the  Latin  diminutive  for  frog,  given  because 
many  of  the  species  grow  in  wet  places. 

The  genus  is  by  far  the  largest  in  the  Ranunculacese,  com- 
prising upwards  of  200  species.  90  of  these  are  natives  or 
naturalized  in  North  America ;  of  those  in  the  trade  in  this 
country  five  are  native  here ;  one  in  the  Canaries ;  five  in  Eu- 
rope, and  two  of  these  also  in  Asia.  Those  cultivated  are  so 
indicated  in  the  following  treatment.  Members  of  the  genus 
are  found  in  mountainous  regions,  and  in  cold  and  temperate 
parts  of  the  globe. 

Perennial  (rarely  annual)  herbs :  leaves  alternate,  simple,  en- 
tire, lobed,  dissected  or  divided  :  flowers  yellow,  white  or  rose ; 
sepals  usually  5,  deciduous  or  marcescent-persistent ;  petals  5 
or  more,  conspicuous  or  minute,  nectar  pit  and  scale  at  base ; 
carpels  many,  i-ovuled :  akenes  generally  flattened,  smooth, 


Davis :   RANUNCULI  OF  NORTH  AMERICA.  463 

papillose  or  spiny,  borne  in  a  head  or  spike ;  styles  minute  or 
elongated. 

In  1886  A.  Gray  wrote  a  revision  of  the  North  American 
Ranunculi  found  north  of  northern  Mexico.  This  was  pub- 
lished in  Proc.  Am.  Acad.  21  :  363-378.  In  Syn.  Flora  I  : 
20-39,  tne  ^vision  is  brought  down  to  1895.  Since  that  date 
the  list  of  species  has  rapidly  increased  and  since  Gray's  first 
revision  two  new  North  American  genera  have  been  segregated 
from  this  one.  In  1892  N.  L.  Britton  discussed  six  species 
"  R.  repens  and  its  Eastern  North  American  allies,"  Trans.  N. 
Y.  Acad.  Sci.  12  :  2-6.  Britton  and  Brown's  111.  Flora  gives 
31  species  in  eastern  United  States  and  Canada.  In  1880  J. 
Freyn  gave  a  long  treatment  of  about  ten  species  in  Flora,  63 : 
179. 

The  present  treatment  includes  96  species,  eighteen  of  which 
are  found  only  in  Mexico  and  south  of  there. 

TENTATIVE  KEY  TO  SPECIES  AND  GROUPS  OF  SPECIES. 

A.  Sepals  and  petals  deciduous  (except  in  77)  ?  petals  yellow  or 
white,  with  nectary  on  the  claw  covered  by  scale;  sepals  5, 
(rarely  only  3  or  4)  ;  petals  5  or  more;  carpels  not  utricular 
when  mature,  usually  somewhat  compressed. — Sec.  EURANUN- 
CULUS,  Gray. 
B.  Leaves,  at  least  some  of  them,  lobed  or  divided. 

C.  Flowers  yellow  (except  some  cultivated  forms  of  31). 
D.  Plants  terrestrial. 

E.   Plants  not  spreading  by  rooting  branches  or  stolons,  ex- 
cept in  12,  26  and  27. 
F.  Sepals  glabrous  or  pubescent  but  not  densely  clothed 

with  black  or  brown  wool. 
G.  Akenes  armed  or   clothed  with  prickles,   spines  or 

prominent  papillae i.   arvensis;2.   muricatus; 

3.  parviflorus ;  4.   hebecarpus ;  5.    Galeottii. 
GG.  Akenes  nearly  smooth  or  pubescent. 

H.   Leaves,  at  least  some  of  the  radical  ones,  divided, 

the  leaflets  either  sessile  or  stalked. 
I.   Radical  leaves  with  some  of  the  leaflets  stalked. 
J.   Petals  short,  about  the  length  of  the  sepals, 
or  shorter. 

K.  Head  of  fruit  globose 6.  alceus. 

KK.  Head  of  fruit  oblong  to  cylindric. 

7.   Pennsylvanicus. 
JJ.  Petals  longer  than  the  sepals. 


464  MINNESOTA    BOTANICAL    STUDIES. 

K.   Beaks  of  akenes  not  hooked. 

L.  Petals  about  5   (or   6),    except   perhaps 

in  13. 

M.   Head  of  fruit  longer  than  thick. 
N.   Sepals    reflexed. 

8.   Macounii;  9.   Sardous. 
NN.  Sepals  spreading  ip.  micranthus; 
\\.fascicularis;  12.  septentrionalis. 
MM.   Head  of  fruit  globose. 

N.  Stem  leaves   present ;    roots  fibrous. 

13.   Hooker i;    14.   pilosus ;    15. 

Bloomeri;    16.  hispidus. 

NN.  Stem   leaves  present,  roots  from 

a  thickened  bulb 17.  bulbosus. 

NNN.   Stem  leaves  wanting. 

1 8 .   Icelandicus. 
LL.  Petals  7  to  16  (double  in  21). 

19.  orthorhynchus ;   20.   dichotomus; 
21.   Llavcenus;  22.   macranthus: 
23.    subalpinus, 
KK.   Beaks  of  akenes  recurved  or  hooked. 

24.  canus ;  25.   amarillo;  26.   repens. 
II.   Radical  leaves  with  the  leaflets  all  sessile. 

27.  palmatus ;  28.  Aschenbornianus ; 

29.  acriformis;  30.    Calif 'ornicus ; 

3 1 .   Asiaticus. 

HH.  Leaves,   at  least  the   radical  ones,  usually  not 
parted  to  the   base,    and    in  some   species   only 
lobed  or  cleft. 
I.   Types    found    native    or    naturalized    north  of 

Mexico. 
J.   Sepals  exceeding  the  petals,  or  sometimes  a 

trifle  shorter,  recurved. 

K.   Beaks  of  akenes  minute,  curved,  or  nearly 
wanting — 32.   abortivus ;  33.  sceleratus ; 

34.   eremogenes. 

KK.  Beaks  of  akenes  nearly  half  the  length  of 
the  body,   recurved... 35.  Allegheniensis ; 
36.  recurvatus ;  37.  Bongardi. 
JJ.  Sepals  decidedly  shorter  than  the  petals. 
K.  Akenes  compressed,  or  flat,  with  firm   or 
indurated  margin. 

L.   Sepals  reflexed 38.   occidentalis; 

39.    Turner  i. 


RANUNCULI    OF    NORTH    AMERICA.  465 

LL.   Sepals  spreading. 

40.   acris;  41.    McCallai. 

KK.  Akenes  turgid  or  lenticular,   marginless. 

L.  Head  of  fruit  oblong  or  cylindraceous. 

42.  pedatijidus;    43.   vicinalis ;  44. 

Eschscholtzii;  45  eximius ; 

46.  saxicola. 

LL.  Head  of  fruit  globose  or  oval. 

47.    Suksdorfii;    48.  avails; 
49.  Arizonicus. 

II.  Types  from   Old  World,   cultivated  here,  not 

naturalized 50.  montanus  ; 

5 1 .   corthiiscefolius. 

III.  Types  found  only  '  in  Mexico  and  southward 
(see  also  var.  of  32). 

'  J.  Plants   with    several    slender    scapose    stems 
bearing  only  bracts  and  terminal  flowers. 

52.  longipedunculatus. 
JJ.  Plants  usually  with  true  stems,  i  to  8  inches 

high 53.   Donianus  ; 

54.  multicaulis ;  55.   Mexicanus. 
JJJ.   Plants  with    true    stems  one  foot  high   or 

more 56.   uncinatus ;  57.  petiolaris. 

HHH.  Leaves  all  2  to  4  times  ternately  parted  or  di- 
vided, divisions  i  line  or  less  in  width ;  flowers  few, 
large;  plants  alpine  or  subalpine,  low,  decumbent 

or  spreading 58.  adoneus ;  59-  triternatus. 

HHHH.  Leaves  all  palmately  or  pedately  lobed  or 
divided;  sepals  nearly  equal  to  the  petals;  plants 
low,  tufted,  arctic  or  alpine. 

60.    Grayi ;  61.  pygmceus. 

HHHHH.  Leaves,  some  of  them,  quite  entire  (ex- 
cept in  62)  ;  others  with  a  few  entire  lobes;  plants 

low  and  glabrous 62.  oxynotus ;  63.  digitatus ; 

64.  glaberrimus. 
FF.   Sepals  densely  clothed  beneath  with  black  or  dark 

brown  wool 65.   Macauleyi ;  66.   nivalis. 

EE.  Plants  spreading  by  slender  creeping  stolons  or  root- 
stocks 67.  natans  ;  68.  hyperboreus ;  69.  Lapponicus. 

DD.   Plants  aquatic  or  amphibious. 

70.   delphinifolius ;  71.   Purshii ;  72.   Missouriensis . 
CC.  Flowers  white  (except  in  a  double  garden  form  of  73). 

73.   aconitifolius ;  74.   Pallasii. 

BB.  Leaves  entire  or  only  denticulate  or  crenulate,  not  lobed,  from 
linear  to  oblong- lanceolate  (or  ovate  in  75)  ;  plants  varying  from 


466  MINNESOTA    BOTANICAL    STUDIES. 

erect  to  ci-eeping  and  rooting  at  the  nodes ;   aquatic  or  in  low 

wet  ground,  or  terrestrial Sec.  FLAMMULA. 

C.  Blades   of    stem    leaves   amplexicaul,   leaves    entire;    flowers 

white 75 .   amplexicaulis. 

CC.   Blades  of  stem  leaves  not  amplexicaul ;  flowers  yellow. 
D.   Stamens  numerous. 

E.  Plant  low  or  erect,   not  spreading  by  slender  creeping 
stems. 

F.   Habitat  aquatic 76.  Lambertianus. 

FF.   Habitat  terrestrial. 

G.   Sepals  and  petals  persistent 77.   arnoglossus. 

GG.  Sepals  and  petals  deciduous. 

H.   Claw  of  petal  i  line  long 78-   unguiculatus. 

HH.   Claw  of  petal  not  nearly  so  long. 

I.  Akenes  beaked. 

J.   Stems  2  to  3  feet  high,  often  rooting  at  lower 

nodes 79-   ambigens. 

JJ.   Stems  much  lower,  not  rooting  at  nodes. 
K.  Mature  fruit  glabi'ous. 

L.   Beak  as  long  as  the  akene  body. 

So.   Madrensis. 

LL.   Beak  much  shorter  than  akene  body. 
M.  Petals  4  to  6  lines  long ;  plants  soli- 
tary, not  much  tufted..8i.  alismcefolius. 
MM.  Petals    about  3   lines    long;    plants 
often  tufted  or  covering  the  ground. 

82.  alismellits. 
KK.  Mature  fruit  villous-pubescent. 

83.  Lemmoni. 

II.  Akenes  beakless ;  styles  deciduous. 

84.   oblongifolius. 

EE.  Plant  spreading  by  slender,  or  fistulous  creeping  stems. 
F.  Lower  leaves  sometimes  cordate ;   flower  stems  ascend- 
ing. 

G.  Margins  of  some  of  the  leaves  slightly  denticulate. 

85.   hydrochar aides. 

GG.  Margins  of  leaves  entire 86.  samolifolius. 

FF.   Lower  leaves  never  cordate. 

G.   Petals  no  longer  than- the  sepals 87.  stolonifer. 

GG.  Petals  nearly  twice  the  length  of  the  sepals. 
H.  Number  of  petals  8  to  10;  plant  glabrous. 

88.  vagans. 

HH.  Number  of  petals  4  to  8;  plants  never  entirely 
glabrous. 


Davis  :    RANUNCULI  OF  NORTH  AMERICA.  4G7 

I.  Stems  filiform  or  nearly  so,  and  usually  rooting 
at   each    node;     peduncles    usually    less   than    2 
inches;  radical  leaves  few 89.  reptans. 

II.  Stems  larger,   at  least  at   the   base,  peduncles 

longer. 

J.   Radical  leaves  not  tufted ;    stems  seldom  as- 
cending  90.  Unalaschensis. 

J  J.  Radical  leaves  tufted ;    stem    somewhat  as- 
cending  91.  microlonchus. 

DD.  Stamens  only  i  to  ten. 

E.   Head  of  fruit  oblong;  stem  leaves  distinctly  petioled. 

92.   trachyspermus. 
EE.  Head  of   fruit  small,  globose;    stem    leaves  sessile  or 

nearly  so 93 .  pusillus. 

AA.  Sepals  and  petals  marcescent-persistent ;   petals  white  or  rose, 
with  ample  nectary  and  imperfect  scale;  carpels  wholly  or  partly 

utricular,  but  compressed  and  broad Sec.  CRYMODES,  Gray. 

B.  Plant  and  sepals  somewhat  pubescent 94.  glacialis. 

BB.   Plant  and  sepals  glabrous. 

C.  Akenes  about  3  lines  long,  wholly  utricular ;   plant  about  6 

inches  high,  stoutish 95.   Andersoni. 

CC.  Akenes  hardly  half  as  large,  not  inflated ;  plant  taller  and 
more  slender 96.  juniperinus. 

1.  R.  arvensis  LINN.  Sp.  PI.  555.     1753. 

Glabrous  or  sparsely  pubescent,  erect,  i  to  2  feet  high, 
branched  above  :  lower  leaves  petioled,  others  sessile  or  nearly 
so,  nearly  all  divided;  leaflets  either  stalked  or  sessile,  cleft  or 
parted  into  linear-oblong  segments  :  petals  yellow,  2  to  3  lines 
long  ;  sepals  of  same  length,  spreading  :  akenes  few,  flattened, 
armed  with  long  spines ;  beak  half  their  length  or  more,  stout ; 
head  depressed-globose.  Europe.  Naturalized  in  New  Jersey, 
and  near  northern  seaports. 

2.  R.  muricatus  LINN.  Sp.  PI.  555.     1753. 

Sparsely  pubescent  or  glabrous,  often  erect,  succulent, 
branched  near  the  base,  6  to  20  inches  high :  lower  leaves  on 
long  broad  petioles,  reniform  to  round  cordate,  3-5~cleft  and 
coarsely  crenate-dentate :  petals  deep  yellow,  3  lines  long ; 
sepals  shorter,  spreading :  akenes  compressed,  large,  con- 
spicuously muricate-spiny ;  beak  stout,  slightly  curved :  head 
loose,  globose.  Asia  and  Europe.  Naturalized  near  towns  in 
Virginia  to  Louisiana,  also  in  California  and  southern  Oregon. 


468  MINNESOTA    BOTANICAL    STUDIES. 

3.  R.  parviflorus  LINN.  Sp.  PI.  2  ed.  780.     1763. 
R.  trachysperimcs  ELL.  Sketch  2:  65.      1824. 

Hairy,  4  to  10  inches  high,  very  slender,  spreading,  branch- 
ing :  leaves  petioled,  reniformto  cordate-orbicular,  y2  to  i  inch 
broad,  3-cleft  or  parted  or  divided,  segments  cuneate,  oval, 
obtuse,  cut  and  toothed  ;  the  upper  leaves  sometimes  5-parted, 
short-petioled :  peduncles  short,  slender ;  petals  pale  yellow, 
hardly  i  line  long ;  sepals  about  the  same  length :  akenes 
oblique,  very  flat,  margined,  papillose ;  beak  short,  sharp. 
Europe.  Naturalized  in  waste  places,  Maryland,  North  Caro- 
lina, Florida,  west  to  Arkansas  and  Texas. 

4.  R.  hebecarpus  HOOK.  &  ARN.  Bot.  Beech.  316.     1841. 
R.  parviflorus  var.  TORR.  &  GRAY,  Fl.  i :  25,  659.    1838. 
R.  hebecarpus  var.  pustllus  WATS.  Bot.  Calif.  I  :  8.     1876. 

Plant  shaggy-hairy,  slender,  ^  to  i  foot  high,  branched: 
leaves  reniform  to  roundish,  small,  3~5-parted  or  divided,  seg- 
ments sessile  or  subsessile,  often  laciniately  cleft :  peduncles 
short ;  petals  hardly  a  line  long,  pale  yellow ;  sepals  about 
equalling  the  petals:  akenes  few,  semi- oval,  compressed, 
clothed  with  recurved  bristles;  beak  short,  subulate,  recurved: 
head  small.  Washington  through  western  California  to  lower 
California. 

5.  R.  Caleottii  TURCZ.  in  Bull.  Soc.  Nat.  Mosc.  27 :  2  :  276. 
1854. 

Roots  not  seen :  plants  otherwise  annual :  stem  somewhat 
branched,  radical  and  lower  stem-leaves  3-parted,  the  divisions 
stalked,  3-lobed  or  parted,  the  lobes  toothed  or  cut,  acute  or 
obtuse,  appressed  pilose ;  petioles  openly  pilose  with  appressed 
hairs  ;  upper  stem-leaves  3-parted,  the  highest  one  sometimes 
not  lobed  :  sepals  reflexed,  openly  pilose;  petals  longer  than 
sepals,  obovate -oblong,  obtuse  :  akenes  compressed,  margined, 
tubercles  on  the  sides,  style  deciduous.  Oaxaca,  Mex.  Alt. 
7,000  to  9,000  feet. 

6.  R.  alceus  GREENE,  Erythea,  3:  69.     1895. 

One  foot  high  or  less,  slender,  branching,  soft-hirsute  and 
villous  :  leaves  about  i  inch  long,  and  much  like  those  of  R. 
canus:  petals  roundly  obovate,  about  i  line  long,  yellow: 
akenes  many,  obliquely  obovoid,  glabrous ;  beak  stout,  re- 
curved :  head  globose.  Elk  mountains,  Mendocino  County, 
Calif. 


Davis :   RANUNCULI  OF  NORTH  AMERICA.  469 

7.  R.  Pennsylvanicus  LINN.  F.  Suppl.  272.     1781. 
R.  Canadcnsis  JACQ^.  Misc.  2:  343.      1781. 

R.  tn'folius  MOENCH.  Meth.  Suppl.  70.     1802. 

R.  hispidus  PURSH,  Fl.  2:  395.     1814.     Not  Michx. 

R.  fascicularis  WATS.  Bot.  King  Exp.  9.      1871. 

Plant  hirsute  or  hispid,  stout,  erect,  8  to  20  inches  high, 
very  leafy,  but  the  radical  leaves  often  dying  down  :  leaves 
petioled,  ternately  compound ;  leaflets  well  stalked,  3-parted 
and  cleft,  much  incised  and  toothed,  segments  acute :  flowers 
small,  yellow,  on  short  peduncles  ;  petals  oblong  to  obovate,  i 
to  2  lines  long ;  sepals  about  the  same  length,  reflexed  :  re- 
ceptacle hairy :  akenes  i  line  long,  oblique  or  semi-oval,  com- 
pressed, roughened ;  beak  subulate,  stout,  short,  nearly 
straight :  head  of  fruit  oblong  to  cylindric.  Wet  ground, 
Nova  Scotia  to  Georgia  west  to  Arizona  and  British  Columbia 
Jacq.  Ic.  Rar.  /.  705. 

8.  R.  Macounii  BRITTON,  Trans.  N.  Y.  Acad.  Sci.   12  :  3. 

1892. 

R.  hispidus  HOOK.  Fl.  I  :    19.      1829.     Not  Michx. 
R.  repens  var.    hispidus  TORR.    &    GRAY,    Fl.    i  :    658. 

1838.     In  part. 

Erect  or  declined,  hairy,  branching,  i  to  2  feet  long,  stems 
rather  few  leaved  :  leaves  ternately  compound,  leaflets  usually  on 
slender  stalks,  crenate,  variously  cleft  and  lobed,  segments 
acute  :  petals  yellow,  obovate,  about  3  lines  long  ;  sepals  shorter, 
often  reflexed,  falling  early  :  akenes  smooth  ;  beak  subulate, 
flat,  short  and  sharp :  head  oblong  or  oval.  Moist  places, 
western  Ontario  to  British  Columbia  south  to  Iowa,  and  in 
mountains  to  Arizona. 

Var.  Oreganus  n.  var. 
R.  hispidus  var.  Oreganus  GRAY,  Proc.  Am.  Acad.  21 : 

376.     1886. 
R.  Oreganus  HOWELL,  Fl.  N.  W.  Am.  i  :   19.      1897. 

Plant  often  taller,  smoothish  or  with  scattered  hairs  :  flowers 
often  larger.  Shaded  wet  grounds,  Willamette  valley,  Ore., 
to  Frazer  valley,  east  to  Kootenai  lake,  Brit.  Col. 

9.  R.  Sardous  CRANTZ,  Stirp.  Austr.  2:  84.     1763. 
R.  parvulus  LINN.  Mant.  1 :   79.     1767. 

R.  Philonotis  EHRH.  Beitr.  2:   145.     1788. 
R.  hirsutus  CURT.  Fl.  Lond.  2:  t.  40.     1821. 


470  MINNESOTA    BOTANICAL    STUDIES. 

Plant  hirsute  especially  below,  3  to  15  inches  high  :  lower 
leaves  3-parted  or  3-foliate,  middle  leaflets  stalked,  others  often 
sessile,  all  obovate-cuneate  to  roundish,  cleft  and  toothed  as  in 
R.  repens :  petals  yellow,  4  to  6  lines  long;  sepals  much 
shorter,  reflexed  :  akenes  flat,  orbicular,  thin-margined ;  beak 
short-subulate :  head  oblong.  Asia  Minor,  northern  Africa, 
Europe.  Naturalized  at  Savannah,  Norfolk,  Philadelphia, 
New  York  and  St.  John,  N.  B. 

10.  R.  micranthus  NUTT.  ex  Torr.  &  Gray,  Fl.  i :   18.    1838. 
R.  abortivus  var.  mtcranthtis  GRAY,  Man.  5  ed.  42.    1867. 

Allied  to  JR.  abortivus,  but  more  slender,  villous  ;  roots  of 
slender  tubers  :  most  of  the  lower  leaves  3-parted,  or  divided 
with  the  leaflets  stalked :  receptacle  glabrous  or  nearly  so. 
April  to  May.  Massachusetts,  New  York  to  Colorado,  and 
Saskatchewan. 

11.  R.  fascicularis  MUHL.  Cat.  54.     1813. 

Roots  a  fascicle  of  thickened  fibres  or  tubers :  plant  finely 
pubescent  throughout,  3  to  10  inches  high,  tufted :  leaves 
mostly  radical,  long-petioled,  3-  (rarely  5-)  divided  ;  middle  leaf- 
let stalked,  others  usually  sessile,  deeply  lobed  and  cleft  into 
oblong  segments  :  petals  5  to  6,  bright  yellow,  obovate-oblong, 
rounded  at  apex,  4  to  6  lines  long ;  sepals  much  shorter, 
spreading  :  akenes  flattened,  slightly  margined,  glabrous  ;  beak 
nearly  their  length ;  head  ovate  or  oblong.  April  to  May. 
Ontario,  New  England  to  Texas,  and  Manitoba.  Meehan's 
Mo.  2.  t.  i.  R.  apricus  GREENE,  Pitt.  4:  145,  1900,  is  a  form 
from  Indian  Territory  with  the  leaflets  rather  narrow  and  some- 
times entire. 

Var.  Deforest!  n.  var. 

Differs  from  the  type  in  having  all  leaves  but  the  first  radical 
ones  cleft  into  linear  to  spatulate  lobes  :  roots  less  thickened  : 
plant  3  to  4  inches  high  :  petals  5  to  10,  linear  to  oblong,  2  to 
4  lines  long.  Collected  by  Harry  P.  DeForest  (G.  42)  near 
Rossville,  111.,  April,  1885. 

12.  R.  septentrionalis  POIR.  in  Lam.  Encycl.  6  :  125.    1804. 
?  R.  lucidus  POIR.  1.  c.  113. 

R.  tomentosus  POIR.  1.  c.  127. 

?  R.  Philonotis  PURSH,  Fl.  2  :  393.      1814. 

R.  JEteZvtsuDC.  Syst.  i:  291.     1818. 


Davis:   RANUNCULI  OF  NORTH  AMERICA.  471 

R.  fascicularis  SCHLECHT.  Animad.  Ranunc.  2  :  30.    t.  2. 

1820.     Not  Muhl. 
R.  SchlechtendaUittoQiL.  Fl.  i :  21.     1829.     (As  to  type.) 

Plant  glabrous  or  sometimes  pubescent,  i  to  3  feet  high, 
branching,  lower  branches  often  rooting  at  their  nodes,  and  run- 
ning some  distance  :  lower  petioles  very  long  :  leaves  composed 
of  3-stalked  leaflets,  which  are  mostly  cuneate  and  cleft  into 
broad  lobes  :  petals  yellow,  obovate,  6  lines  long ;  sepals  half 
as  long,  spreading  :  akenes  much  compressed,  widely  margined  ; 
beak  nearly  as  long,  subulate,  flat:  head  of  fruit  rather  small, 
ovoid.  Often  in  low,  wet  places.  New  Brunswick  to  Georgia, 
northwestward  to  Winnipeg. 

13.  R.  Hookeri  SCHLECHT.  in  Linnaea,  9:  610.     1835. 

Stem  erect,  branched,  i  to  2  feet  high :  lower  leaves  and 
petioles  pilose  with  spreading  yellowish  hairs,  upper  leaves 
with  peduncles  sulcate,  pubescence  appressed  :  leaves  subpin- 
nate,  nearly  as  narrow  as  in  R.  repens :  sepals  reflexed ;  petals 
narrow ;  receptacle  pilose :  akenes  oblique,  roundly  obovate, 
laterally  margined,  marked  with  minute  impressions  and  often 
a  few  scattered  tubercles  on  sides,  acuminate ;  base  of  style  per- 
sistent :  head  of  fruit  globose.  Allied  to  /?.  acris  in  habit  and 
roots ;  to  R.  repens  in  foliage.  Its  narrow  often  numerous 
petals  and  reflexed  calyx  remove  it  from  all.  Vera  Cruz, 
Oaxaca,  in  San  Miquelito  Mts.,  and  other  places  in  Mexico. 
Common  in  grass  lands. 

14.  R.  pilosus  H.B.K.  Nov.  Gen.  &  Sp.  5 :  36.     1821. 

Roots  thick-fibrous,  many :  stem  ascending,  somewhat  di- 
chotomously  branched  above,  hirsute  :  radical  leaves  petiolate, 
ternate,  appressed-pilose,  10  to  15  lines  wide,  i  to  i^  inches 
long  ;  leaflets  cut-toothed,  lateral  ones  sessile,  ovate-rhomboid, 
sometimes  2-3-lobed ;  terminal  leaflets  large,  on  stalks  2  to  6 
lines  long,  subrotund,  3-lobed  or  3-parted ;  lower  stem  leaf  like 
the  radical  leaves  but  much  dissected,  short-petioled ;  upper 
stem  leaves  gradually  less  dissected,  those  near  the  flowers 
sessile :  flowers  as  large  as  in  R.  bulbostis,  peduncles  silky ; 
sepals  5,  ovate,  acutish,  pubescent  outside,  reflexed,  much 
shorter  than  the  petals,  deciduous ;  petals  5  (rarely  6),  oblong, 
with  rounded  tips,  5  lines  long,  glabrous,  supplied  with  a  scale 
on  the  claw :  akenes  glabrous,  oblong  to  obovate,  compressed, 


472  MINNESOTA    BOTANICAL    STUDIES. 

tapering  into  the  persistent  style ;  fruit  in  globose  head.     High 
altitudes,  Guatemala.     Also  near  Bogota,  U.  S.  of  Colombia. 

15.  R.  Bloomeri  WATS.  Bot.  Calif.  2:  426.     1880. 

R.  Chilensis  HOOK.  &  ARN.  Bot.  Beech.  134.  1841. 
Stem  ascending,  i  to  2  feet  long,  sparsely  hairy  or  becoming 
glabrous :  radical  leaves  bright  green,  long-petioled,  some 
broadly  cordate  or  ovate,  coarsely  dentate  or  incised,  others  3- 
parted,  some  divided  into  3  leaflets  which  are  short-stalked  and 
the  middle  one  often  3-lobed  ;  stem  leaves  short  petioled  :  petals 
yellow,  6  lines  long,  emarginate ;  sepals  shorter :  akenes  gla- 
brous, 2  lines  long,  turgid ;  beak  slender,  subulate,  persistent. 
San  Francisco  bay. 

16.  R.  hispidus  MICHX.  Fl.  i:  321.     1803. 

R.  Marilandicus  POIR.  in  Lam.  Encycl.  6:  126.      1804. 

R.  repens  var.  Marilandicus  TORR.   &  GRAY,  Fl.  1:21. 
1838. 

R.  fascicularis  BRITTON,  PI.  N.  J.  3.  1881. 
Appressed-pubescent,  when  young  densely  villous :  stems 
slender,  ascending  or  spreading,  ^  to  2  feet  long :  leaves  pal- 
mately  3-parted,  or  pedately  or  pinnately  3~5~divided ;  the  di- 
visions ovate,  or  variable,  middle  one  often  stalked,  others 
usually  sessile  ;  all  often  cuneate  at  base,  sharply  cleft  or  lobed  : 
petals  5  or  more,  light  yellow,  3  to  6  lines  long  ;  sepals  half  as 
long,  spreading :  akenes  broadly  oval,  lenticular,  margined,  ab- 
ruptly tipped  ;  beak  half  their  length,  subulate,  slightly  curved  ; 
head  ovoid  to  globose.  Earliest  spring.  Canada  through 
Eastern  and  Middle  States  to  Florida  and  Arkansas. 

17.  R.  bulbosus  LINN.  Sp.  PI.  554.     1753. 

R.  speciosus  HORT.    ex  VILM.   Fl.   PL   Terre   i  ed.  722. 

1865. 

Plant  from  a  true  bulb,  erect,  about  i  foot  high,  hairy :  leaves 
petioled,  3~5-parted,  the  divisions  sometimes  stalked,  segments 
lobed  :  flowers  terminating  the  branches,  bright  yellow,  large ; 
petals  large,  obovate,  shining  above  ;  sepals  much  smaller,  often 
reflexed :  akenes  compressed,  with  short  beak,  borne  in  a  glo- 
bose head.  Spring  and  summer.  Persia,  Europe,  northern 
Africa.  The  double  form  is  perhaps  best  suited  for  culture. 

18.  R.  Icelandicus  n.  sp. 

Caudex  short,  roots  fibrous  :  plant  pubescent  throughout :   no 


Da-sis:    RANUNCULI  OF  NORTH  AMERICA.  473 

true  stem,  scape  about  3  inches  high,  nearly  erect,  slender: 
leaves  about  i  inch  long  on  petioles  the  same  length,  blade  3- 
divided  or  parted,  the  leaflets  sessile  or  the  middle  one  stalked  ; 
segments  with  about  3  entire  or  toothed  cuneate  lobes  :  petals  5, 
yellow,  large,  obovate-cuneate,  obcordate  or  retuse ;  sepals 
shorter,  spreading,  pubescent :  carpels  much  like  those  of  R. 
acris.  Collected  June,  1895,  by  Elizabeth  Taylor  at  Seydis- 
fjordr,  Iceland. 

19.  R.  orthorhynchus  HOOK,  ex  GRAY,  Proc.  Am.  Acad.  21 : 

377-     1887- 
R.  ornithorhynchus  WALP.  Rep.  1 :  43.      1842  (by  error). 

Root  thick  fibrous  :  plant  10  to  18  inches  high,  erect,  branched, 
hirsute  to  nearly  glabrous  :  leaves  oblong  in  outline,  pinnately 
compound  ;  5  to  7  leaflets  cleft  and  incised,  quite  variable  ;  up- 
der  leaflets  often  confluent  and  sessile  or  nearly  so,  lower  ones 
well  stalked  :  petals  7  to  16,  yellow,  rarely  purple  beneath,  ob- 
ovate,  4  to  6  lines  long  ;  sepals  much  shorter,  pubescent  beneath, 
reflexed,  deciduous :  akenes  glabrous,  obliquely  ovoid,  com- 
pressed, i  to  2  lines  long,  margined ;  style  of  same  length, 
straight,  rigid,  persistent :  head  globose.  May  to  July.  Wet 
places.  British  Columbia  to  western  Oregon  and  Montana. 
Cultivated. 

Var.    platyphyllus  GRAY,  Proc.  Am.  Acad.  21 :  377.    1886. 
R.  macranthus  WATS.   Bot.    King   Exp.  9.     1871.     Not 

Scheele. 
R,  maximus  GREENE,  Bull.  Torr.  Club,  14:   118.      1887. 

Often  3  feet  or  more  high  :  leaves  larger,  2  to  4  inches  across, 
the  leaflets  often  3  inches  long,  and  laciniately  cut :  petals  often 
larger  than  the  type.     Wasatch  Mountains,  northern  Utah,  Py 
ramid  lake,  northern  Nevada,  northern  California,  Washington, 
Idaho.     Cultivated. 

20.  R.  dichotomus  Moc.  &  SESSE.   ex   DC.   Syst.   i :  288. 
1818. 

Stem  erect,  often  dichotomously  branched :  radical  leaves 
very  long-petioled,  bipinnate  :  flowers  yellow ;  sepals  reflexed  : 
akenes  with  acuminate  erect  beaks.  Mexico. 

21.  R.  Llavsenus  SCHLECHT.  in  Linnaea,  10 :  233.     1836. 
Stem  prostrate,  the  flowers  on  erect  or  ascending  branches, 

terminal :  leaves  3-divided   and   again   3-lobed,  segments  nar- 


474  MINNESOTA    BOTANICAL    STUDIES. 

rowly  and  sharply  cut-toothed ;  middle  leaflet  stalked,  lateral 
ones  sessile,  all  cuneate  at  base  ;  the  long  petioles  hirsute,  widely 
sheathing  and  smooth  at  the  base  :  peduncles  sulcate  ;  recepta- 
cle pilose  :  flowers  yellow ;  calyx  reflexed :  akenes  obliquely 
round-obovate,  slightly  margined  laterally,  terminated  by  the 
long  erect  style,  smooth  :  head  of  fruit  globose.  June  to  July. 
Meadows,  Jalapa,  Vera  Cruz.  Allied  to  R.  dichotomus,  leaves 
nearly  as  finely  dissected,  yet  much  allied  to  R,  Hookeri. 

22.  R.  macranthus  SCHEELE,  in  Linnaea,  21 :  585.     1848. 
R.  repens\ai-.  macranthus  GRAY,  PL  Lindh.  2  :  141.    1850. 

Plant  hairy ;  erect  or  spreading,  y2  to  3  feet  high  :  leaves  3- 
5-divided,  the  middle  leaflet  longer  stalked  than  the  others,  lobed 
and  cleft  into  narrower  segments  than  in  R.  septentrionalis : 
petals  7  to  16,  yellow,  5  to  7  lines  or  longer,  oblong  toobovate; 
sepals  shorter,  spreading :  receptacle  hairy :  akenes  flat,  ovate 
to  orbicular,  widely  margined;  style  subulate,  long,  often  only 
partly  persistent:  head  large,  slightly  lengthened.  Texas, 
southwestern  Arizona  into  Sonora,  Mex. 

23.  R.  subalpinus  n.  n. 

R.  detyhinifolius  H.B.K.  Nov.  Gen.  &  Sp.  5  :  38.      1821. 
Not  Torr. 

Roots  fibrous :  stem  erect,  branched  above,  few-flowered, 
silky-hairy :  radical  leaves  long-petioled,  pilose  on  both  sides, 
ternate,  lateral  leaflets  subsessile,  2-parted,  terminal  one  well 
stalked,  3-parted,  segments  2-3-lobed,  incised  and  toothed ; 
lobes  lanceolate  ;  stem  leaves  similar  but  smaller,  short-petioled  : 
flowers  on  long  peduncles,  erect,  as  large  as  in  R.  bulbosus; 
sepals  5,  silky  outside,  reflexed,  ovate,  acutish,  yellowish,  much 
shorter  than  the  corolla ;  petals  about  15  (Jide  BonpL),  yellow, 
glabrous,  5  lines  long,  spatulate-oblong,  apex  rounded,  claw 
furnished  with  a  scale :  young  ovaries  many,  small,  sessile, 
ovate  to  subrotund,  compressed,  glabrous ;  style  long  and  slen- 
der. May.  Moist  places.  Altitude  8,000  to  9,000  feet.  San 
Miguelito  Mountains  and  at  Guanajuato,  Mex. 

24.  R.  canus  BENTH.  PL  Hartw.  294.     1848. 

R.  Californicus  var.  canus  WATS.  Bot.  Calif.  I  :  8.     1876. 

Plant  canescent  when  young  but  often  becoming  green  and 

sparingly  villous ;  erect  or  ascending,  rather  large,  i  to  2  feet 

high :  leaves  with  mostly  3  or  more  divisions  ;  the  middle  one 


Davis:   RANUNCULI  OF  NORTH  AMERICA.  475 

stalked  ;  leaflets  cuneate,  2— 3-cleft  and  again  incised :  petals 
yellow,  5  to  6  lines  long,  obovate  ;  sepals  half  as  long,  reflexed, 
soft-hairy  :  akenes  about  2^  lines  long  ;  beak  less  than  half  as 
long,  broad,  hooked.  Sacramento  valley,  Calif. 

Var.  Blankinshlpii  ROBINSON,  Syn.  Fl.  i :   i  :  35.     1895. 

The  silky  coat  persistent  but  less  dense  than  in  the  typical 
plants :  akenes  plainly  hispid  and  papillose.  Capay,  Yolo 
County,  Calif. 

Var.  hesperoxys  n.  var. 
R.  hesperoxys  GREENE,  Erythea,  2  :  189.     1894. 

Plants  much  greener  than  the  type ;  doubtless  due  to  the 
early  falling  of  the  canescence.  California. 

25.  R.  amarillo  BERTOL.  Fl.  Guat.  24.     1840. 

Hirsute,  stem  branched,  ascending  :  lower  leaves  petioled  ; 
leaves  compound  ;  leaflets  stalked,  subcordate-ovate,  acute,  3- 
lobed,  cut-toothed ;  the  upper  leaves  often  short-petioled,  ter- 
nate,  divisions  lanceolate,  dentate  :  petals  about  8,  yellow,  ob- 
long-cuneate ;  sepals  shorter,  hairy,  reflexed ;  flowers  as  large 
as  R.  acris :  akenes  compressed,  glabrous  ;  style  long,  erect  but 
recurved  at  tip  :  head  of  fruit  globose.  Guatemala. 

26.  R.  repens  LINN.  Sp.  PI.  554.     1753. 

R.  pro  stratus  POIR.  in  Lam.  Encycl.  6:   113.     1804. 
R.  Clintonii  BECK.  Bot.  9.     1833. 

Roots  fibrous :  plant  more  or  less  hairy :  spreading  by 
runners;  flower  stems  often  ascending  6  to  12  inches:  leaves 
petioled,  3-divided  ;  middle  leaflet  or  all  of  them  stalked,  often 
again  3-lobed  or  cleft,  and  somewhat  coarse  toothed,  bases 
cuneate  or  truncate  :  petals  obovate,  5  to  6  lines  long  ;  sepals 
much  shorter,  spreading,  hairy  below :  akenes  compressed, 
margined;  beak  short,  stout,  slightly  bent:  head  globose. 
May  to  July.  Common.  Nova  Scotia  and  Newfoundland  to 
Virginia,  westward.  Also  Europe  and  Asia.  Cultivated. 

27.  R.  palmatus  ELL.  Sketch,  2:  61.     1824. 

Included  by  Gray,  '86,  with  R.  septentrionalis  which  it  is 
much  like;  plant  smaller,  more  decumbent;  runners  often 
long :  leaves  i  inch  across,  thin,  somewhat  3-parted  or  divided, 
divisions  ovate,  coarsely  few-toothed ;  lowest  leaves  often  sub- 
entire  :  flowers  6  lines  broad.  Pine  lands  and  swampy  places, 
Tennessee,  South  Carolina,  Florida. 


476  MINNESOTA    BOTANICAL    STUDIES. 

28.  R.  Aschenbornianus  SCHAU.  in  Linnasa,  20  :  719.     1847. 
Stem    erect,     many-flowered :     leaves    hairy    to    subhirsute ; 

radical  leaves  long-petioled,  ternate  to  bipinnate,  the  pinnules 
3-parted  to  many  lobed,  the  lobes  nearly  linear;  peduncles 
silky :  sepals  reflexed :  receptacle  subpilose :  akenes  com 
pressed,  slightly  margined,  smooth,  with  fine  impressions  or 
punctures,  style  straight :  fruit  in  a  globose  head.  Mountains 
of  Mexico  near  "  Tutam." 

29.  R.  acriformis  GRAY,  Proc.  Am.  Acad.  21 :  374.     1886. 
R.  acris  HOOK.  Fl.  i:   18.      1829  (partly). 

Plant  with  short  rather  appressed  pubescence,  slender,  erect, 
i  foot  or  more  in  height :  leaves  all  3~7-parted  or  divided ; 
divisions  2-3-cleft  or  lobed,  into  lanceolate  or  linear  segments 
which  are  often  entire :  petals  yellow,  roundly  obovate,  about 
3  lines  long ;  sepals  about  half  as  long,  spreading  or  becoming 
reflexed:  akene  i  to  2  lines  long:  beak  half  as  long,  curved. 
Eastern  Rockies  in  Alberta  ;  Montana,  Wyoming  ;  wet  places. 
Southern  Colorado  at  10,000  feet. 

R.  Montanensis  Rydb.  Mem.  N.  Y.  Bot.  Gard.  i  :  166, 
1900,  is  a  form  with  beak  more  slender  and  more  curved. 

30.  R.  Californicus  BENTH.  PI.  Hartw.  295.     1848. 

R.  acris  var.  Deppi'i  NUTT.  ex  Torr.  &  Gray,  Fl.  i  :   21. 

1838. 
R.    delphinifolius   TORR.    &    GRAY,    Fl.   i :  659.      1838. 

Not  Torr. 
R.    dissectus    HOOK.    &.    ARN.   Bot.    Beech.  316.     1841. 

Not  Bieb. 
R.  regulosus  GREENE,  Pitt.  2  :  58.      1890. 

Roots  fibrous:  plant  rather  weak,  y2  to  2  feet  high,  usually 
pubescent  or  hirsute ;  branching  and  without  leaves  in  upper 
part:  leaves  ternately  divided  or  parted,  or  palmately  5-divided 
into  linear  or  narrow  often  2-3-parted  divisions  :  petals  6  to  15, 
glossy  yellow,  oblong  or  narrowly  obovate,  4  to  6  lines  long  : 
akenes  flat,  slightly  margined,  nearly  2  lines  long :  beak  very 
short.  Rather  dry  places.  Western  California  and  adjacent 
Oregon,  common.  Cultivated. 

Var.  Ludovicianus  (GREENE). 

R.  Ludovicianus   GREENE,    Bull.    Cal.  Acad.  Sci.  2  :  58. 
Mch.  6      1886. 


Davis:    RANUNCULI  OF  NORTH  AMERICA.  477 

7?.  Californicus  var.  latilobtis  GRAY,  Proc.  Am.  Acad.  21  : 
375.  1886. 

Basal  leaves  3-parted,  divisions  broadly  or  narrowly  cuneate, 
incisely  cleft  or  laciniate  :  stem  leaves  not  so  much  dissected. 
Southern  California. 

Var.  crassifolius  GREENE,  Erythea,  1 :   125.     1893. 

Stout  and  low,  sparingly  villous  throughout :  lower  leaves 
not  so  deeply  parted  as  the  type,  coarsely  toothed  ;  stem  leaves 
mostly  deeply  parted  into  3  oval  or  oblong  quite  entire  segments  : 
flowers  and  akenes  larger  than  in  the  type.  Ft.  Bragg,  Men- 
docino  Co.,  Calif. 

31.  R.  Asiaticus  Linn.  Sp.  PI.  552.     1753. 

Roots  fleshy  :  plant  erect,  either  simple  or  branched,  ^  to  i 
foot  high  :  leaves  petiolate,  becoming  sessile  toward  the  top, 
ternate  or  biternate  ;  segments  toothed  or  deeply  3-lobed  ;  flow- 
ers terminating  in  the  stems  and  branches,  variable  in  color : 
calyx  spreading,  becoming  reflexed ;  petals  large,  obovate, 
blunt :  fruits  in  spike.  May  to  June.  Asia  Minor.  Flor.  des 
Serr.  16:  1679  (fl-pl).  Revue  Hort.  Belg.  1890:  133  (var. 
superbissimus).  Sibth.  Fl.  Gr.  518.  The  cultivated  forms  of 
this  species  are  constantly  increasing  in  number.  They  are  of 
two  main  types  :  (i)  The  florist's  section  called  Persian  Ranun- 
culi or  true  R.  Asiaticus.  (2)  The  gardener's  section,  called 
Pivoine  and  Turban  Ranunculi,  or  var.  Africanus.  There  are 
many  named  forms  of  each  in  the  American  trade. 

32.  R.  abortivus  LINN.  Sp.  PL  551.     1753. 
R.  nitidus  WALT.  Car.  159.      1788. 

Sparingly  pubescent  or  quite  glabrous,  one-half  to  2  feet 
high,  branched :  stem  leaves  sessile  or  short-petioled,  once  or 
twice  3-parted  or  lobed,  segments  oblong  or  linear,  somewhat 
cuneate ;  lower  leaves  long-petioled,  lucid  green,  crenate  or 
lobed,  broadly  cordate,  roundish,  or  ovate:  petals  pale  yellow, 
hardly  over  a  line  long ;  sepals  longer  and  larger,  reflexed ; 
receptacle  short,  pubescent :  akenes  compressed,  glabrous, 
tipped  with  the  minute,  curved  beak :  head  small,  globose. 
Spring.  Moist  grounds  and  woods.  Labrador  to  Florida, 
north  and  west  to  Colorado  and  British  Columbia.  Var.  ency- 
clus  FERNALD,  Rhodora,  1:52,  1899,  a  slender,  flexuose  form 
with  thinner,  glossy,  orbicular,  radical  leaves. 


478  MINNESOTA    BOTANICAL    STUDIES. 

Var.  Harveyi  GRAY,  Proc.  Am.  Acad.  21 :  372.     1886. 
R.  abortivus  var.  grandiflorus  ENGELM.  ex  Branner  &  Co- 

ville,  Ark.  Geol.  Surv.  6:   162.     1891. 
R.  Harveyi  BRITTON,  Mem.  Torr.  Club,  5:   159.     1894. 
Stem  and  foliage  more  slender  and  roots  often  thicker  than 
in  the  type  :   plant  somewhat  pubescent :    petals  of  the  extreme 
forms   3   lines   long  and   much  longer  than  the  sepals  :   akenes 
sometimes  few,  large   and   in  a  globose  head,  but  varying  to 
those  of  the  type.     Damp  rocks,  Arkansas  to  St.  Louis,  Mo. 
Var.  australis  BRAND.  Zoe  4  :  399.     1894. 
Lower  leaves   large,  reniform :    petals    2    to    3    lines   long ; 
flowering  in  August.     Abundant  in  wet  places,  high  summits 
of    Sierra  de   la   Laguna   and   San  Fransisquito,  Lower  Cali- 
fornia. 

33.  R.  sceleratus  LINN.  Sp.  PI.  551.     1753. 

Stems  stout,  hollow  :  plant  glabrous  or  nearly  so,  one-half  to 
2  feet  high,  branching :  radical  and  lower  stem  leaves  thick, 
long-petioled,  3~5-lobed,  reniform  or  cordate ;  lobes  cuneate, 
crenately  incised  or  cleft;  upper  stem  leaves  sessile  or  petioled, 
deeply  lobed  or  parted  ;  the  lobes  cuneate-oblong  or  linear, 
toothed  or  entire  :  petals  i  to  2  lines  long,  yellow ;  sepals  often 
a  little  shorter :  akenes  numerous,  very  small,  compressed, 
glabrous,  barely  apiculate  :  head  oblong.  April  to  Aug.  Wet 
ditches  and  shallow  water.  New  Brunswick  to  Florida  and 
somewhat  westward.  Europe  and  Asia. 

34.  R.  eremogenes  GREENE,  Erythea,  4 :  121.     1896. 

R.  sceleratus  var.  multijidus  NUTT.  ex  Torr.  &  Gray,  Fl. 
I  :  19.  1838. 

Leaves  more  dissected  than  R.  sceleratus:  stem  nearly  leaf- 
less :  head  of  akenes  nearly  globose  or  ovate,  large.  Habitat 
of  that  species,  eastern  base  of  Rockies  in  Colorado  to  the 
Sierra  Nevadas,  to  northwest  British  America  and  south  to 
Arizona. 

Var.  degener  GREENE,  Pitt.  4:  144.     1900. 

Stems  several,  short,  ascending  :  roots  coarser  than  in  the 
type :  akenes  with  no  marginal  development :  head  more 
rounded.  Southern  Colorado. 

35.  R.  Allegheniensis  BRITTON,  Bull.  Torr.  Club,  22:  224. 
1895. 

R.  abortivus  HOOK.  Fl.  1 :  15.     1829.     In  part. 


Davis:   RANUNCULI  OF  NORTH  AMERICA.  479 

This  is  also  closely  allied  to  R.  abortivus  in  habit  and  foliage. 
Plant  glabrous,  not  lucid :  akenes  slightly  compressed  and 
margined,  tipped  with  subulate  hooked  or  recurved  styles  hardly 
half  their  length.  April,  May,  North  Carolina,  Virginia, 
Massachusetts. 

36.  R.  recurvatus  Pom.  in  Lam.  Encycl.  6:  125.     1804. 
R.  lanuginosus  WALT.  Car.  159.      1788.     Not  Linn. 
R.  saniculceformis  MUHL.  Cat.  54.      1813. 

R.  tomentosus  SPRENG.  Neue  Entd.  i  :  287.     1820. 

Plant  hirsute  or  only  slightly  hairy,  erect,  ^  to  2  feet  high, 
branching :  all  the  leaves  petioled  and  never  divided  to  the 
base,  i^  to  3  inches  wide,  deeply  3-cleft,  the  lobes  broadly 
cuneate,  acute,  toothed  or  lobed  :  petals  light  yellow,  about  2 
lines  long ;  sepals  of  same  length  or  little  longer :  akenes  com- 
pressed, margined  :  beak  one-half  their  length,  recurved.  Damp 
woods,  Nova  Scotia  to  Lake  of  the  Woods,  south  to  Missouri 
and  Florida. 

37.  R.  Bongardi  GREENE,  Erythea,  3:  54.     1895. 

R.  occidentals  var.  parviflorus  TORR.  Bot.  Wilkes  Exp. 

214.     1854. 
R.  occidentals  var.  Lyalli  GRAY,  Proc.  Am.  Acad.  21 : 

373.     1886.     Not  R.  Lyalli  Hook.  f. 
R.  tencllus  var.  Lyalli  ROBINSON,  Syn.  Fl.  I :  i  :  33.  1895. 
R.  Greener  HOWELL,  Fl.  N.  W.  Am.  i:   18.     1897. 
R.  Earlei  GREENE,  Pitt.  4  :   15.     1899. 
R.  Lyalli  RYDB.  Mem.  N.  Y.  Bot.  Gard.  i :  166.     1900. 

Much  like  the  following  variety  which  is  better  known  and 
was  formerly  considered  the  type.  Plant  more  hirsute  :  leaf 
segments  much  broader :  petals  rather  of  the  larger  size,  some- 
what persistent :  akenes  somewhat  hispid ;  styles  rather  long. 
Northern  California  to  Colorada  and  Montana  and  to  Fort 
Wrangel,  Alaska. 

Var.  Douglasii  (HOWELL). 
R.  recurvatus  BONG.   Veg.    Sitch.   123.     1831.     Mainly, 

not  Poir. 
R.  tenellus    NUTT.   ex    Torr.  &   Gray  Fl.   i  :   23.     1838. 

Not  Viviani. 

R.  Nelsoniv&v.  tenellus  GRAY,  Proc.  Am.  Acad.  8:  374. 
1872. 


480  MIXXESOTA    BOTAXICAL    STUDIES. 

R.  occidentalis  var.  tencllus  GRAY,  Proc.  Am.  Acad.  21 : 

373-     1886. 

7?.  occidentalis  var.  Eiseni  GRAY,  1.  c.  (in  small  part). 
7?.  Nehoni  var.   glabriusculus    HOLZINGER,    Cont.    Nat. 

Herb.  3:  210.      1895.     Not  7?.  glabriusculus  Rupr. 
R.  Bongardi 'var.  tenelhis  GREENE,  Erythea,  3  :  54.    1895. 
R.  Douglasn  HOWELL,  Fl.  N.  W.  Am.  i  :   18.     Mch.  15. 

1897. 
R.  arcuatus  HELLER,  Bull.  Torr.  Club,  24:  310.    June  29. 

1897. 

Slender,  erect,  usually  over  one  foot  high,  slightly  pubescent 
or  glabrous,  hirsute  on  petioles  and  peduncles  :  leaves  deeply 
3-5-cleft ;  segments  broadly  cuneate  to  oblanceolate,  coarsely 
toothed  :  petals  i  to  2  lines  long  ;  receptacle  glabrous  :  akenes 
compressed,  glabrous,  semi-oval ;  styles  persistent,  circinate- 
revolute ;  head  small,  globose.  Southern  California,  Idaho,  to 
Alaska. 

38.  R.  occidentalis  NUTT.  ex  Torr.  &  Gray,  Fl.  i :  22.    1838. 
R.  recurvatus  var.  Nehoni  DC.  Syst.   i:   190.      1818. 
R.  recurvatus  (2  forms)  SCHLECHT.  Animad.   Ranunc.  2  : 

28.      1820. 

R.  Schlechtendalii  HOOK.  Fl.  i  :   21.    1829.    (As  to  plant.) 
R.  Nehoni  GRAY,  Proc.  Am.  Acad.  8:  374.      1872. 
R.  Eiseni  KELLOGG,    Proc.    Calif.    Acad.    Sci.    7:     115. 

1877. 
R.  occidentalis  var.  Eiseni  GRAY,  Proc.  Am.  Acad.   21  : 

373.      1886  (mainly). 
7?.  occidentalis  \ax.  brevistylus  GREENE,  Pitt.  3  :  14.     1896. 

Plant  villous  with  spreading  hairs,  5  to  18  inches  high  :  lower 
leaves  petioled,  round  cordate,  3~5-cleft  or  parted ;  segments 
cuneate-obovate,  often  2-3-cleft  and  cut ;  some  leaves  3-divided 
with  the  leaflets  stalked  ;  upper  stem  leaves  smaller  with  lanceo- 
late segments  :  petals  yellow,  large,  spreading ;  sepals  half  as 
long,  reflexed  :  akenes  glabrous  or  sparsely  bristled  :  style  flat- 
tened, subulate,  hooked,  half  as  long  as  akenes;  receptacle  gla- 
brous :  head  of  fruit  ovoid.  Low  open  places.  Alaska  to 
Montana  and  California. 

Var.  Howellii  GREENE,  Pitt.  3  :   14.     1896. 
/?.  Ho-wellii  GREENE,  ex  Howell,  Fl.  N.  W.  Am.  i :   17. 
1897. 


Davis:   RANUNCULI  OF  NORTH  AMERICA.  481 

Rather  leafy ;  upper  leaves  more  deeply  and  repeatedly 
cleft :  styles  longer,  slender,  subulate,  nearly  straight.  Dry 
hills,  Ashland,  Oregon,  southward  toward  Klamath  river, 
Calif. 

Var.  ultramontanus  GREENE,  Pitt.  3:   13.     1896. 

Plant  tufted ;  divisions  of  lower  leaves  not  cuneiform,  deeply 
cleft  into  lanceolate  segments  ;  upper  ones  lanceolate,  entire  : 
several  flowers  ^  inch  across  :  styles  hooked.  Moist  places, 
Truckee  river,  east  of  Sierras,  Calif. 

Var.  Rattani  GRAY,  Proc.  Am.  Acad.  21  :  373.     1886. 
R.  Rattani  HOWELL,  Fl.  N.  W.  Am.  1 :   17.     1897. 
R.  ciliosus  HOWELL,  1.  c. 

Differs  from  the  type  in  having  the  akenes  covered  with 
short,  stiff  hairs  and  also  roughened  with  papillae.  Josephine 
county,  Ore.,  to  central  California. 

Var.  robustus  GRAY,  Proc.  Am.  Acad.  21 :  373.     1886. 
R.  occidentalis  GRAY,  Proc.  Am.  Acad.  8:  374.      1872. 

Stem  stout,  often  a  foot  high  :  flowers  large,  9  to  15  lines 
long,  long-peduncled  :  petals  broadly  obovate  :  akenes  numer- 
ous, very  large.  Unalaska  and  islands  westward. 

39.  R.  Turner!  GREENE,  Pitt.  2  :  296.     1892,  except  syn. 

Plant  appearing  much  like  a  tall  specimen  of  R.  occidentalis : 
petals  longer,  4  to  7  lines  long  :  flowers  long-peduncled  :  more 
akenes  in  a  head  ;  styles  circinately-revolute,  strongly  so.  Por- 
cupine river,  Alaska. 

40.  R.  acris  LINN.  Sp.  PI.  554.     1753. 

Plant  hairy  up  to  the  sepals,  erect,  ^  to  3  feet  high,  often 
branched  :  radical  leaves  on  long  slender  petioles  ;  others  with 
shorter  petioles  sheathing  the  stem,  or  nearly  sessile  :  leaves  3- 
parted  nearly  to  the  base,  the  divisions  ovoid-cuneate,  2-3-lobed 
and  coarsely  toothed  or  cut ;  bracts  linear,  lobed  or  entire : 
flowers  yellow,  9  to  12  lines  across,  several,  on  rather  short  pe- 
duncles ;  sepals  hairy  beneath,  ovate,  shorter  than  the  petals  ; 
petals  5,  glabrous,  obovoid,  obtuse,  bearing  a  prominent  scale 
at  base  :  akenes  compressed,  coriaceous  on  margins ;  style  very 
short :  head  globose.  May  to  September.  Newfoundland, 
Canada,  Eastern  States.  Said  to  be  naturalized  from  Europe. 
Var.  Jlore-pleno  HORT.  is  more  used  in  the  trade.  Bot.  Mag. 
215- 


482  MINNESOTA    BOTANICAL    STUDIES. 

41.  R.  McCallai  n.  sp. 

Stem  erect,  slender,  12  to  20  inches  high,  branched  toward 
the  top,  somewhat  pubescent :  radical  leaves  hairy,  on  slender 
hairy  peduncles  2  to  4  inches  long ;  blade  3~5-parted  nearly  or 
quite  to  the  base,  segments  less  than  i  inch  long,  cuneate,  di- 
vergent, cleft  into  2  to  3  linear  lobes  ;  stem  leaves  none  or  bract- 
like,  subtending  the  branches,  or  a  small  one  near  the  middle  : 
flowers  2  to  6,  large,  yellow  ;  petals  5,  obovate,  entire  or  obcor- 
date ;  sepals  shorter,  spreading,  hairy :  carpels  ovate  in  a  glo- 
bose head ;  styles  subulate,  hooked  :  receptacle  glabrous.  Col- 
lected by  W.  A.  McCalla  (2113)  near  Banff,  Alberta,  Canada, 
July,  1899,  in  wet  meadows.  Differs  from  R.  acris  in  its 
leaves,  usually  naked  stem,  etc. 

42.  R.  pedatifidus  J.   E.    SMITH   in    Rees'    Cycl.   No.    72. 
1813-16. 

R.  arcticus  RICHARDS,  in   Frankl.   ist  Journ.  i   ed.  App. 

741.      1823. 

R.  affinis  R.  BR.  Parry  ist  Voy.  App.  265.      1824. 
R.  amcenus  LEDEB.  Fl.  Alt.  2:  320.      1830. 
R.  auricomus  HOOK.  f.  Arc.  PL  283,  312.     1862. 

Plant  sparsely  hairy,  3  to  15  inches  high,  slender,  sometimes 
branched :  radical  and  lower  stem  leaves  petioled,  broadly 
ovate,  crenate,  toothed,  lobed,  or  cleft  nearly  to  the  base  into 
segments  which  are  often  narrow  ;  upper  stem  leaves  deeply 
cleft,  nearly  sessile,  lobes  narrow  :  petals  yellow,  3  to  4  lines 
long ;  sepals  shorter,  pubescent  beneath  :  akenes  often  hairy? 
with  short  beak  :  head  oblong  to  cylindric.  Qjiebec  to  Arctic 
regions,  west  to  Alaska ;  Rockies  to  Colorado  and  Arizona. 
Ledeb.  Ic.  /.  nj. 

Var.   cardiophyllus   BRITTON,  Bull.  Torr.   Club,  18 :    265. 

1891. 

R.  cardiophyllus   HOOK.  Fl.  I  :   14,  and  vars.  t.  6.     1829. 
R.  affinis  var.  lasiocarpus  TORR.  Bot.  Wilkes   Exp.  213. 

1854. 
R.  affinis  var.  leiocar-pus  TRAUT.  ex    Midden.   Reise  in 

Sibir.  62.      1856. 
R.  affinis  var.  cardiophyllus  GRAY,  Proc.  Am.  Acad.  Phil. 

1863:  56. 

R.  affinis  var.  validus   GRAY,  Proc.  Am.  Acad.  21 :  371. 
1886. 


Davis:    RANUNCULI  OF   NORTH  AMERICA.  483 

R.  ajfinis  var.  micropetalus  GREENE,  Pitt.  2:   no.     1890. 
R.  Arizonicus  var.  subajfinis  GREENE,  1.  c.  60.    Not  Gray. 
R.  inamcenus  GREENE,  Pitt.  3:  91.      1896. 
Differs  from  the  type  in  its  stouter  habit,  radical  leaves  often 
cordate   at  base,  usually  not  much  lobed,  but  variable  :  flowers 
larger :    akenes   either  hairy  or  glabrous.     New   Mexico   and 
Arizona  to  Montana,  east  to  Labrador. 
Var.  pinetorum  (GREENE). 
R.  cardiophyllus  var.  pinetorum   GREENE,   Pitt.  4:    144. 

1900. 

Stem  short,  canescently  villous  :  roots  strongly  and  copiously 
developed  :  leaves  oval,  often  subcordate,  or  truncate  at  base, 
margins  crenate :  flower  i  inch  across  :  head  of  akenes  not  so 
long[as  in  the  type,  ovoid  or  globose.  Pine  woods,  Graham's 
Park,  southern  Colorado.  7,800  feet. 

43.  R.  vicinalis  GREENE,  Pitt.  4:   145.     1900. 

This  is  an  Alaskan  plant  from  the  region  of  Fort  Selkirk, 
which  differs  from  R.  pedatifidus  in  its  larger  flowers,  and  in 
having  the  radical  leaves  cleft  or  parted  into  about  7  lobes,  and 
these  again  3-cleft.  But  we  find  this  leaf  character  in  some 
Colorado  forms  of  that  type. 

44.  R.  Eschscholtzii  SCHLECHT.     Animad.  Ranunc.   2  :   16. 
/.  /.     1819. 

R.  nivalis  var.  Eschscholtzii  WATS.   Bot.  King  Exp.  8. 
1871. 

R.  ocreatus  GREENE,  Pitt.  4:  15.  1899. 
Caudex  short,  oblique,  roots  fibrous  :  plant  slightly  hairy,  3-8 
inches  high  :  leaves  roundish  or  broader  in  outline  ;  stem  leaves 
3~5-lobed  or  parted,  lobes  lanceolate  to  oblong  or  linear 
spatulate  ;  basal  leaves  with  broader  lobes,  or  lobed  like  the 
others  :  flowers  i  to  3  ;  petals  yellow,  broadly  obovate,  some- 
times slightly  crenulate  or  obcordate,  3  to  6  lines  long ;  sepals 
pubescent  beneath :  akenes  lenticular,  margined,  glabrous ; 
beak  sharp,  straight  or  sometimes  recurved :  head  of  fruit  ob- 
long or  nearly  globose.  North  Alaska  in  mountains  to  south- 
ern California,  east  to  Colorado. 

45.  R.  eximius  GREENE,  Erythea,  3:   19.     1895. 

R.  alpeophilus  A.    NELSON,    Bull.   Torr.    Club,    26:   350. 

1899. 
Much  like  R.  Eschscholtzii,  but  often  larger,  nearly  glabrous  : 


484  MINNESOTA    BOTANICAL    STUDIES. 

radical  leaves  broader,  less  divided  ;  upper  leaves  with  lobes 
1-2  inches  long  :  sepals  nearly  glabrous  :  akenes  broadly  oval 
or  obovate.  Mountains  of  Wyoming,  Idaho,  Colorado. 

46.  R.  saxicolaRvDB.,  Mem.  N.  Y.  Bot.  Gard.  i :  164.    1900. 
Allied  to  R.   Suksdorfii;  differs   slightly  in  the  form  of  the 

lower  leaves,  which  are  often  more  reniform-flabellate  :  akenes 
pubescent :  head  of  fruit  oblong.  Cedar  mountain  and  Mill 
Creek,  Montana,  Yellowstone  Park. 

47.  R.  Suksdorfii  GRAY,  Proc.  Am.  Acad.  21 :  371.      1886. 
Roots  fibrous :  stems  slender,  3  to  6  inches  high,  glabrous  : 

radical  and  lowest  stem  leaves  small,  about  6  to  8  lines  long, 
subreniform  to  broadly  flabelliform  with  truncate  base,  deeply 
3~5-cleft  or  parted ;  divisons  cuneate,  again  3~5-cleft  or  in- 
cised;  upper  stem  leaves  with  linear  divisions:  flowers  i  to  3, 
deep  yellow ;  petals  round  obovate,  retuse,  4  to  6  lines  long : 
akenes  turgid-lenticular,  sharp-edged,  glabrous  ;  style  persistent 
for  a  time,  slender,  ^  line  long,  equalling  the  akene  body : 
head  of  fruit,  globular.  July  to  Aug.  Damp  places,  6,000  to 
8,000  feet  alt.  Olympic  mountains,  Mt.  Rainier,  Mt.  Adams 
east  to  the  Blue  mountains,  Oregon  and  into  Montana.  Cul- 
tivated. 

48.  R.  ovalis  RAF.  Proc.  Dec.  36.     1814. 

R.  rhomboideus  GOLDIE,  Edin.  Phil.  Journ.  6:  329.  t.  n. 

f.  i.     1822. 

R.  brevicaulis  HOOK.  Fl.  i  :   13.  t.  7.     1829. 
R.  auricomus  var.    Cassubicus  E.   MEYER   PI.   Labr.   96. 

1830. 

Pubescent,  3  to  15  inches  high:  radical  and  lower  leaves 
roundish  to  ovate-oblong,  crenate,  or  slightly  lobed,  base  trun- 
cate or  cuneate,  petioled,  3~7-divided,  lobes  linear  or  oblong : 
petals  yellow,  narrow,  3  to  6  lines  long ;  sepals  much  shorter : 
akenes  oval,  minutely  beaked:  head  of  fruit  globose.  Wis- 
consin and  northern  Illinois,  north  to  Labrador  and  the  North- 
west Territory. 

49.  R.  Arizonicus  LEMMON  ex  Gray,  Proc.  Am.  Acad.  21 : 
370.     1886. 

R.  affinis  TORR,  Bot.  Mex.  Bound.  29.      1858.     (In  part.) 

Roots  fascicled,  somewhat  thickened  :  plant  glabrous  or  with 

soft-villous  hairs    below,  6  to   12  inches  high:  radical   leaves 


Davis:    RANUNCULI  OF  NORTH  AMERICA.  485 

oval  to  oblong,  cordate,  crenate-dentate ;  the  later  ones  often  5- 
cleft  and  again  3-5-lobed  ;  stem  leaves  i-3-ternate  with  narrow 
linear  divisions  :  petals  yellow,  often  6  to  7,  ovate  to  oblong,  3 
to  5  lines  long :  akenes  compressed,  thin-margined,  pubescent : 
head  small,  globose.  Willow  Spring,  and  mountains  of  south- 
ern Arizona. 

Var.  subaffinis  GRAY,  Proc.  Am.  Acad.  21 :  370.     1886. 

R.  Arizonicus  GREENE,  Pitt.  2  :  60.      1890. 

7?.    subsagittatus    var.    subaffinis    GREENE,    Pitt.  2:    no. 
1890. 

R.  subaffinis  RYDB.  Bull.  Torr.  Club,  24:  246.     1897. 

Plant  lower  than  the  type,  usually  i-flowered  :  akenes  densely 
pubescent,  with  subulate  style  nearly  their  own  length :  head  of 
fruit  oval.  High  altitudes.  Mt.  Agassiz,  in  San  Francisco, 
mountains  of  Arizona.  Also  Chihuahua. 

Var.  subsagittatus  GRAY,  Proc.  Am.  Acad.  21 :  370.    1886. 

Stouter  than  the  above ;  villous  at  least  at  first :  stem  simple, 
few-flowered :  radical  leaves  thick,  oblong,  bases  subcordate  to 
sagittate :  petals  broader  than  in  the  type :  head  of  akenes 
larger,  oval.  Wet  ground.  Northern  Arizona  to  San  Fran- 
cisco mountains. 

50.  R.  montanus  WILLD.  Sp.  PL  2:  1321.     1799. 

Rootstock  creeping,  1-3  inches  long,  %  inch  thick :  plant  6 
inches  high,  pubescent  with  soft  appressed  or  spreading  hairs, 
especially  toward  the  top  :  radical  leaves  few,  petiolate,  smooth, 
orbicular  in  outline,  3-parted,  and  lobed  into  blunt,  toothed 
segments ;  stem  leaves  sessile  or  nearly  so,  clasping  the  stem, 
3~5-parted  into  narrow  somewhat  toothed  or  entire  lobes  :  flow- 
ers solitary,  terminating  the  simple  or  once-branched  stem,  i 
inch  or  larger ;  sepals  concave,  acute,  yellowish-green,  slightly 
hairy;  petals  5,  large,  broadly  obovoid,  bright  yellow,  with 
small  scale  and  pore  at  base :  akenes  turgid,  glabrous ;  beak 
strongly  hooked,  puberulent.  May  to  July.  Europe.  Culti- 
vated. Bot.  Mag.  3022.  Bot.  Cab.  1610. 

Var.  dentatus  BAUMG.  Enum.  Stirp.  Magn.  Trans.  2:   124. 

About  1823. 
R.  carpaticus  Herbich.*  Sel.  PL  Rar.  Galic  15.     1836. 

Leaves  much  more  toothed  than  in  the  type :  plant  much 
taller:  flowers  larger.  Cultivated.  Bot.  Mag.  7266.  Garden 
52:  1138- 


486  MINNESOTA    BOTANICAL    STUDIES. 

51.  1809.    R.   corthusaefolius  WILLD.   Enum.   Hort.   Berol. 
588. 

Root  of  thick,  fleshy,  fasciculated  fibers  :  plant  velvety  hairy, 
i  to  3  feet  high  :  lower  leaves  long-petioled,  roundish  to  reni- 
form,  incised,  and  with  cut  and  toothed  lobes ;  stem  leaves 
divided  into  3  to  5  narrow  lobes  ;  upper  ones  sessile :  flowers 
several  or  many,  terminal  and  axillary,  rather  paniculate  ;  sepals 
5,  ovate  to  lanceolate,  green  with  pale  margins  ;  petals  5,  large, 
broadly  obovate,  glossy  yellow :  akenes  compressed,  hairy  on 
sides,  tapering  into  recurved  styles  nearly  their  own  length  : 
head  of  fruit  short  oval.  May,  Island  of  Teneriffe,  Canary 
group.  Garden,  45  :  944.  Bot.  Mag.  4625.  Not  very  hardy, 
and  needs  protection  in  winter  and  early  spring.  It  is  well 
suited  for  pot  culture.  It  is  increased  by  division  of  the  roots  in 
autumn. 

52.  R.  longipedunculatus   SCHEIDW.  in  Hortic.    Beige.    5: 
163.  t.  9.     1838. 

Roots  fibrous :  stem  slender,  weak,  scapose,  often  once 
branched  near  the  base  :  plant  pilose  with  yellowish  hairs  :  rad- 
ical leaves  3-lobed,  middle  lobe  trifid,  otherwise  mostly  entire ; 
true  stem  leaves  wanting  or  very  low ;  petioles  i  to  i  ^  inches 
long  :  peduncles  4  to  5  inches  long,  slender,  erect  or  ascending, 
bibracteate  :  flowers  i  or  2,  terminal;  petals  12  to  15,  oblong- 
lanceolate,  acutish,  yellow  ;  sepals  reflexed.  Wet  places,  Real 
del  Monte,  north  of  Mexico  City. 

53.  R.  Donianus  PRITZ.  ex  Walp.  Rep.  2:  740.     1843. 

R.  humilis  D.  DON,  ex  G.  Don,  Gen.  Syst.  i :  34.     1831. 
Not  Pers.  1807. 

Short,  i  to  3  inches  high,  pilose  :  radical  leaves  stalked,  cor- 
date, obtuse,  slightly  3-lobed  and  crenate  :  peduncles  long,  rad- 
ical, axillary  and  terminal :  flowers  small,  yellow :  carpels 
rather  inflated,  beaked.  Mexico. 

54.  R.  multicaulis  D.  DON,  ex  G.  Don,  Gen.  Syst.  i :  34. 
1831. 

Plant  about  3  to  4  inches  high,  pilose  :  stems  numerous,  pros- 
trate or  ascending  :  radical  leaves  p'etioled,  cordate-roundish,  3- 
lobed ;  lobes  crenate :  stem  leaves  sessile,  entire,  opposite  or 
apparently  so  :  flowers  yellow,  medium  size  ;  petals  emarginate, 
much  longer  than  the  sepals  which  are  reflexed  :  carpels  rather 
inflated,  pointed  :  head  of  fruit  ovate.  Mexico. 


Davis:    RANUNCULI  OF  NORTH   AMERICA,  487 

55.  R.  Mexicanus  n.  n. 

R.  gcoides    H.B.K.   Nov.   Gen.  and  Sp.   5:    37.   t.  429. 
1821.     Not  Siev.  1794. 

Roots  fibrous  :  stem  5  to  8  inches  high,  simple  or  branched 
low ;  plant  silky  pubescent  up  to  and  including  under  side  of 
sepals :  radical  leaves  3-parted  nearly  to  midrib :  divisions 
toothed  or  incised,  ovate-cuneate ;  petiole  i  to  2  inches  long  ; 
stem  leaf  with  3  to  5  narrow  segments,  sessile  or  on  petiole 
sheathing  the  stem ;  bracts  linear :  flowers  (rarely  2  to  3), 
large,  terminating  long,  erect  peduncles  ;  sepals  5,  ovate,  much 
shorter  than  petals,  reflexed  ;  petals  10,  spatulate  oblong,  rather 
obtuse,  5-7  lines  long,  spreading,  claw  provided  with  nectar 
pit  and  scale  :  akenes  compressed,  smooth ;  style  as  long  as 
akene  body,  persistent  for  a  time,  erect.  May.  Real  del 
Monte,  north  of  Mexico  City;  Guajuco,  Nuevo  Leon,  northern 
Mexico. 

56.  R.   uncinatus    D.  DON  ex  G.  Don,   Gen.    Syst.   i :    35. 
1831. 

Roots  fibrous  :  stem  erect,  i  foot  high  :  plant  glabrous  :  rad- 
ical leaves  not  seen  ;  stem  leaves  long-petioled,  3-parted,  seg- 
ments 3-lobed ;  lobes  toothed  or  again  lobed,  acute;  leaves 
near  the  flower  ternate,  leaflets  linear-lanceolate,  acute,  quite 
entire :  flowers  small,  yellow,  terminal  and  lateral,  on  slender 
peduncles :  akenes  few,  ending  in  hooked  beaks :  head 
globose.  Mexico. 

57.  R.  petiolaris  H.B.K.  Nov.  Gen.  and  Sp.  5:  36.  t.  428. 
1821. 

Roots  fibrous :  stem  erect,  striate,  puberulent,  somewhat 
branched  above,  about  6-flowered  :  radical  leaves  on  petioles  6 
to  7  inches  long,  3-parted  nearly  to  the  midrib ;  the  divisions 
often  2-3-lobed  and  incised ;  the  lobes  often  somewhat  oblong- 
cuneate,  deep  green  and  appressed-pubescent  above,  pale  and 
appressed-pilose  beneath ;  stem  leaves  short-petioled,  smaller 
and  with  much  more  slender  lobes ;  bracts  linear-lanceolate, 
lobed  or  entire ;  base  of  petioles  membranaceous,  somewhat 
sheathing  :  flowers  medium  size,  on  erect  peduncles  which  are 
pilose;  sepals  5,  ovate-oblong,  acute,  pilose  beneath,  shorter 
than  the  petals,  reflexed,  deciduous;  petals  5,  obovate,  rounded 
at  the  end,  a  claw  and  scale  at  base,  yellow,  glabrous,  about  4 


488  MINNESOTA    BOTANICAL    STUDIES. 

lines  long,  much  exceeding  the  stamens  :  akenes  oblique,  com- 
pressed, glabrous,  rather  abruptly  joined  to  the  short,  persistent 
style:  head  of  fruit  subglobose.  September.  Near  "Los 
Joares" ;  and  Santa  Rosa,  state  of  Mexico.  Altitude,  8,400  feet. 

58.  R.  adoneus  GRAY,  Proc.  Acad.  Phila.  1863:  56. 

JR.  amcenus  GRAY,  Am.  Journ.  Sci.  Ser.  2,  33:  241.   1862. 

Not  Ledeb. 
R.  orthorynchus  var.   alpinus  WATS.  Bot.  King  Exp.  9. 

1871. 

Root  slender- fibrous  :  plant  shaggy-hairy,  4  to  12  inches  high, 
sometimes  becoming  decumbent :  leaves  usually  2-3-times  3- 
parted  and  lobed,  lobes  all  narrow-linear,  acute;  primary  di- 
visions of  leaves  sessile  or  nearly  so ;  petioles  of  basal  leaves 
membranous  in  lower  part ;  stem  leaves  sessile  or  on  a  sheath- 
ing base,  usually  borne  opposite  resembling  an  involucre  :  petals 

5  (or  6  to  8),  large,  yellow,  rounded  outwardly,  cuneate  at  base, 

6  lines  long,  much  exceeding  the  lanceolate   sepals  which  are 
hairy  beneath :    akenes  somewhat  compressed,    acutish :  style 
long,  straight,  subulate ;  head  globular  to   oblong.     Summer. 
Rockies  of  Colorado.     Altitude  10,000  feet.     Cultivated.     In- 
troduced 1881. 

59.  R.  triternatus  GRAY,  Proc.  Am.  Acad.  21 :  370.     1886. 

Roots  fascicled,  fleshy-fibrous :  plant  low :  leaves  often  3- 
times  3-divided  and  parted ;  leaflets  long-petioled,  their  lobes 
narrow-linear  to  linear-spatulate  and  obtuse  :  petals  yellow,  4  to 
5  lines  long,  obovate :  akenes  turgid,  not  margined ;  beaks 
slender  :  receptacles  thick  :  head  of  fruit  globose.  Near  Golden- 
dale,  S.  Wash. 

60.  R.  Grayi  BRITTON,  Bull.  Torr.  Club,  18 :  265.     1891. 
R.  -pedatifidus  HOOK.  Fl.  1 :   18.  /.  18.     1829.     Not  Smith. 
R.  Hooker i  REGEL.  Bull.    Soc.   Nat.    Mosc.    34  :   2  :  47. 

1861.     NotSchlecht. 
R.  Drummondi  GREENE,  Erythea,  2:   192.     1894. 

Rather  stout,  i-2-flowered :  basal  leaves  either  biternately 
or  pedately  divided  and  parted  into  linear  oblong  or  spatulate 
lobes,  main  divisions  often  stalked :  stem  leaves  similar,  only  i 
or  2  :  petals  3  lines  long ;  sepals  shorter,  sparsely  and  finely 
villous  :  akenes  each  about  i  line  long,  borne  in  a  globular  head. 
Lat.  52°  to  55°,  on  eastern  Rockies,  Gray's  Peak,  Colo.,  and 
near  Ironton,  12,000  to  13,000  feet. 


Davis :    RANUNCULI  OF  NORTH  AMERICA.  489 

61.  R.  pygmseus  WAHL.  Fl.  Lapp.  157.  /.  8.  f.  i,     1812. 
R.  Lap-ponicus  OED.  Fl.  Dan.  t.  14.4..     1762.     Not  Linn. 

Very  minute,  i  or  2  inches  high,  puberulent  or  glabrous  : 
leaves  3~5-lobed  or  divided,  2  to  5  lines  wide,  lower  ones  on 
slender  petioles,  others  subsessile  :  flowers  2  to  3  lines  across  ; 
petals  yellow,  little  longer  than  the  sepals  :  akenes  lenticular ; 
beak  slender:  head  of  fruit  somewhat  oblong,  2  lines  long. 
High  Rockies  of  Montana  to  Colorado,  polar  regions  across 
America,  Greenland,  Europe  and  Asia. 

Var.  Sabinii  n.  var. 
JR.  Sabinii  R.  BR.  Parry  istVoy.  App.  264.     1824. 

Flowers  larger  than  the  type  :  sepals  hairy.     Montana. 

62.  R.   oxynotus    GRAY,   Proc.  Am.  Acad.  10 :  68.     1874. 

Caudex  short,  roots  fibrous  :  plant  glabrous,  4  to  10  inches 
high  :  radical  leaves  in  a  numerous  tuft,  6  to  10  lines  across, 
mostly  round-reniform,  with  several  roundish  lobes  or  deep  cre- 
nations  :  stem  leaves  i  or  2,  flabelliform  to  cuneate,  3~5-cleft  or 
parted  ;  lobes  lanceolate-linear  to  oblong  :  petals  yellow,  broadly 
obovate,  4  to  5  lines  long,  exceeding  the  sepals  :  akenes  com- 
pressed, semiovate,  glabrous,  about  i  line  long;  beak  strong, 
subulate  :  head  of  fruit  6  lines  long  :  receptacle  thick  and  fleshy. 
Mineral  King  Mt.,  Mariposa  Co.,  and  central  Sierras,  all  in 
California. 

63.  R.  digitatus  HOOK.  Kewjourn.  3:   124.  /.  4..     1851. 
Very  low,  glabrous  :  roots  a  cluster  of  slender  tubers  :  stem 

leaves  few,  subsessile,  2-4-parted ;  lobes  oblong-lanceolate  to 
nearly  linear ;  radical  leaves  similar  or  entire  and  lanceolate, 
petiolate  :  petals  5  to  n,  yellow,  spatulate-oblong,  3  to  5  lines 
long  :  akenes  slightly  compressed  and  margined  ;  styles  slender  : 
head  very  small,  often  elongated.  Yellowstone  Park  to  north- 
ern Nevada. 

64.  R.  glaberrimus  HOOK.  Fl.  i :  12.  t.  5.     1829. 

R.  brevicaulis  HOOK.  Lond.  Journ.  Bot.  6:  66.     1847. 

R.  Austina  GREENE,  Erythea,  3:  44.  1895. 
Root  a  cluster  of  thickened  fibers  :  plant  rather  succulent,  4 
to  10  inches  high,  glabrous  :  radical  leaves  roundish  to  oblan- 
ceolate  or  spatulate  ;  base  tapering  or  obtuse,  often  2-5-lobed 
above,  or  crenate  or  entire ;  stem  leaves  usually  deeply  3-lobed 
or  parted,  lobes  entire  :  petals  yellow,  broadly  obovate,  3  to  6 


490  MINNESOTA    BOTANICAL     STUDIES. 

lines  long  ;  sepals  nearly  as  long,  often  purple  beneath :  akenes 
puberulent  or  glabrous,  lenticulate,  slightly  margined,  with 
small,  short  beak  :  fruit  in  a  globose  to  oblong  head.  Early 
spring.  British  Columbia  to  California  and  Colorado. 

Var.  ellipticus  GREENE,  Fl.  Francis,  298.     1892. 
R.  ellipticus  GREENE,  Pitt.  2  :  no.     1890.     3:92.      1896. 

Basal  leaves  elliptic-lanceolate  to  oblong,  entire  or  only  once 
lobed  on  one  side  :  petals  often  much  narrower  than  in  the  type  : 
head  of  akenes  drooping  to  the  ground.  Distributed  with  the 
type  in  its  lower  altitudes  and  southern  range. 

65.  R.  Macauleyi  GRAY,  Proc.  Am.  Acad.  15:  45.     1880. 
R.  nivalis  REP.  Chief  Eng.  U.  S.  A.   1878:   1833.     Not 

Linn. 

Some  of  the  roots  thick  and  fleshy :  plant  3  to  7  inches  high ; 
stem  villous-hairy  to  glabrous,  young  leaves  very  villous  on 
margins  :  leaves  thick,  lanceolate  to  ovate-spatulate,  entire  ex- 
cept toward  the  apex,  there  often  coarsely  or  finely  2  to  10 
toothed ;  lower  leaves  petiolate,  others  sessile  or  on  short, 
sheathing  petioles  :  petals  obovate  to  flabelliform,  crenulate,  5 
to  7  lines  long,  yellow ;  sepals  shorter,  densely  coated  beneath 
with  dark  brown  hairs;  peduncles  hairy  :  akenes  smooth,  some- 
what compressed,  slightly  margined ;  styles  linear-subulate, 
persistent,  nearly  straight :  head  of  fruit  ovate  to  oblong  or 
cylindric.  Near  snow  line,  11,500  feet  altitude  in  La  Plata 
mountains  and  San  Juan  Co.,  Colo. 

66.  R.  nivalis  LINN.  Sp.  PI.  553.     1753. 

R.  sulphureus  SOLAND.  in  Phipp's  Voy.  202.      1774. 

A  short  caudex  with  slender  roots :  plant  pubescent  or  be- 
coming glabrous  below,  3  to  7  inches  high  :  lower  leaves  cune- 
ate-flabelliform  to  reniform,  about  3-lobed  or  deeply  cleft ;  lobes 
sometimes  notched ;  upper  leaves  subsessile,  about  5-lobed  or 
parted,  divisions  linear-oblong,  entire :  petals  yellow,  obovate 
to  roundish,  sometimes  emarginate,  3  to  4  lines  long;  sepals 
shorter,  densely  wooly :  akenes  rather  turgid ;  beak  subulate. 
Greenland,  Hudson  Bay  region,  Alaska,  Hall  island,  Behring 
Sea,  south  in  Rockies  to  Lat.  55°,  Northern  Asia  and  Europe. 

67.  R.  natans  C.  A.  MEYER  ex  Ledeb.  Ic.  t.  114.     1830. 
R.  radicans  C.  A.  MEYER  ex  Ledeb.  Ic.  t.  116.     1830. 
R.  Purshii  TORR.    Ann.    Lye.    N.    Y.    2:    162.      1828. 
Not  Richards. 


Davis:   RANUNCULI  OF  NORTH  AMERICA.  491 

R.  hyperboreus  var.  natans  REGEL,  Bull.  Soc.  Nat.  Mosc. 

34:  pt.  2:  43.      1861. 

Much  like  R.  hyperboreus,  but  differing  in  having  leaves 
larger,  reniform  or  truncate  at  base,  lobes  3  to  5,  often  more 
rounded  :  petals  much  larger  ;  receptacle  thickened  and  fleshy  : 
head  of  fruit  larger.  Creeping  and  rooting  in  mud  or  some- 
times floating  in  shallow  water.  Rockies  of  Colorado.  Also 
northern  Asia. 

68.  R.  hyperboreus  ROTTB.  Skrift.  Kjoeb.  Selsk.  10 :  458. 
t.  ./.  f.  16.     1770. 

Low  creeping  plant  with  slender  and  glabrous  stems  and  pet- 
ioles :  one  or  two  leaves  from  each  (rooting)  node,  broadly 
ovate  with  rounded  or  truncate  bases,  3-lobed  or  slightly  cleft, 
margins  of  lobes  nearly  entire  ;  petioles  sheathing  at  the  base  : 
flowers  minute,  few,  yellow  ;  petals  about  equalling  the  reflexed 
sepals  ;  peduncles  i  inch  or  less  in  length  :  akenes  hardly  com- 
pressed ;  beak  almost  wanting  :  head  of  fruit  globose,  hardly  2 
lines  broad.  Wet  soil.  Greenland,  Labrador,  Arctic  Alaska  ; 
also  Europe  and  Asia. 

69.  R.  Lapponicus  LINN.  Sp.  PL  553.     1753. 
Anemone  nudicdulis  GRAY,  in  Bot.  Gaz.  n  :  17.      1886. 

Scapose  from  filiform  rootstocks,  3  to  6  inches  high :  radical 
leaves  long-petioled,  3-parted,  the  divisions  obovate-cuneate, 
obtuse,  crenate  or  lobed :  scape  slender,  taller  than  the  leaves, 
often  with  a  lobed,  bract-like  leaf :  flower  solitary,  yellow ; 
petals  5  to  6 ;  sepals  of  about  the  same  length,  reflexed :  akenes 
a  line  or  more  long,  ovate,  tapering  into  the  persistent,  hooked 
beak.  North  shore  of  Lake  Superior,  west  to  the  Rockies, 
north  to  Arctic  America ;  also  Europe  and  Siberia. 

70.  R.  delphinifolius  TORR.  ex  Eaton,  Man.  2  ed.  395.   1818. 
R.  multifidus  PURSH,  Fl.  2  :  736.     1814.     Not  Forsk. 

R.  flaviatilis  BIGEL.  Fl.  Bost.   i  ed.  1 :  39.     1814.     Not 

Willd. 
R.  lacustris  BECK  &  TRACY,  N.  Y.  Med.  &  Phys.  Journ. 

2  :  112.      1823. 

R.  BeckiiG.  DON,  Gen.  Syst.  i :  39.     1831. 
R.  Purshitv&r.  aquatilis  LEDEB.  Fl.  Ross,  i  :  35.     1841. 
R.  multifidus  var.  terrestris  GRAY,  Man.  5  ed.  41.     1867. 

Aquatic  or  partly  emersed,  with  long  fistulous  stems  :  sub- 


492  MINNESOTA    BOTANICAL    STUDIES. 

mersed  leaves  ternately  decompound  into  narrow  filiform  or 
capillary  divisions,  flaccid,  petioles  very  short  and  sheathing ; 
emersed  leaves  smaller,  and  much  less  dissected,  often  only 
3~7-parted  into  cuneate  lobes,  petioles  often  longer  than  blades; 
young  leaves  from  nodes  taking  root  on  muddy  banks,  still  less 
lobed  and  divided,  their  petioles  and  under  sides  hairy  :  petals 
deep  yellow,  5  to  8,  broadly  obovate,  4  to  6  lines  long,  exceed- 
ing the  sepals  :  akenes  rather  turgid,  obliquely  ovate,  hardly  i 
line  long,  becoming  callous-margined  on  base  and  ventral  edge  ; 
beak  half  their  length,  straight,  compressed :  head  of  fruit  glo- 
bose or  oblong.  Quiet  water,  and  muddy  ditches  and  banks. 
North  Carolina  to  northern  Canada,  west  to  British  Columbia 
and  California.  Also  Siberia. 

71.  R.  Purshii  RICHARDS,  Frankl.  ist  Journ.  741.     1823. 
R.  pusillus  LEDEB.  Mem.  Acad.  Petrop.  5:   546.      1812. 
R.  Gmclini  DC.  Syst.  1 :  303.      1818. 

R.  Langsdorfii  DC.  Prod.  1 :  34.      1824. 

R.  limosus  NUTT.  ex  Torr.  &  Gray,  Fl.  I  :   20.      1838. 

R.  radicans  REGEL,  Bull.  Soc.  Nat.  Mosc.  34 :  pt.  2 : 
44-5.  1 86 1.  Not  Mey. 

R.  multifidus  var.  limosus  LAWSON,  Rev.  Canad.  Ranunc. 
47.  1870. 

R.  multifidus  var.  repens  WATS.  Bot.  King  Exp.  8.  1871. 
Stems  slender,  rooting  at  lower  nodes  and  creeping,  in  muddy 
places,  pubescent  on  younger  parts :  leaves  slender-petioled, 
3  to  12  lines  broad,  palmately  divided  into  obtuse  lobes  and  seg- 
ments :  petals  yellow  or  whitish,  i  to  3  lines  long,  ovate  ;  sepals 
smaller,  falling  early :  akenes  smooth,  y2  line  long,  no  cal- 
loused margin  ;  style  persistent,  slender,  shorter  than  the  body  : 
head  of  fruit  smaller  than  in  R.  del-phinifolius ,  Bogs,  ditches, 
etc.  Arctic  America  to  Northern  Michigan,  west  to  British 
Columbia  and  Washington,  south  to  New  Mexico. 

72.  R.  Missouriensis  GREENE,  Erythea,  3:  20.     1895. 
Much  like  R.  Purshii  in  habit   and  leaves  :  differs  in  being 

sparingly  pubescent :  leaves  wider  than  long,  i  to  3  inches 
wide  :  head  of  fruit  more  oblong  :  akenes  prominently  callous- 
margined  up  one  edge,  sides  wrinkled ;  style  subulate,  y2  as 
long  as  body.  Missouri  to  New  Mexico. 

73.  R.  aconitifolius  LINN.  Sp.  PI.  551.     1753. 

Plant   pubescent,   y2   to  3  feet  high,  branched :  leaves  pal- 


Davis:    RANUNCULI  OF  NORTH  AMERICA.  493 

mately  3~5-parted,  parts  cut-toothed,  upper  ones  sessile  and 
with  oblong  to  linear-lanceolate  lobes  :  flowers  white,  several  on 
a  stem;  sepals  flat,  pubescent;  petals  oblong,  cuneate  to  orbic- 
ular. May  to  June.  Mountains  of  middle  Europe.  Var.Jlore- 
pleno  HORT.,  called  White  Bachelor's  Button,  and  Fair  Maids 
of  France,  has  very  ornamental,  double,  white,  globose  flowers. 
Garden  45,  p.  29  and  48,  p.  506.  Var.  luteus-plcno  HORT., 
flowers  much  doubled  but  of  a  golden  yellow  color.  The  type 
and  varieties  are  used  in  borders  and  half  wild  places. 

74.  R.  Pallasii  SCHLECHT.  Animad.  Ranunc.   i:   15.  t.  2. 
.  1819. 

Plant  creeping,  glabrous  :  stems  and  petioles  large,  hollow ; 
ascending  part  of  stem  naked  or  i-leaved  :  leaf-blades  short, 
linear  to  oblong,  rather  obtuse,  entire  or  sometimes  2-3-lobed  : 
petals  8  to  u,  oblong  to  obovate,  white,  4  to  6  lines  long; 
sepals  3  to  4,  shorter,  greenish,  broad  :  akenesthin-crustaceous, 
2  lines  long ;  beak  short.  In  shallow  water.  Arctic  Alaska, 
St.  Lawrence  islands,  etc.,  across  to  northern  Asia,  and  Lapland. 

75.  R.  amplexicaulis  LINN.  Sp.  PL  549.     1753. 

Stems  erect,  5  to  10  inches  high,  with  two  or  three  flowering 
branches,  glabrous  :  leaves  entire,  ovate  to  lanceolate,  amplex- 
icaul,  acuminate,  glabrous  or  at  first  with  hairy  edges  soon  be- 
coming glabrous,  glaucous  :  flowers  3  to  6,  either  terminal  or 
axillary,  pure  white  with  yellow  stamens ;  sepals  pointed ; 
petals  much  larger,  obtuse.  Mountains  of  southeastern  Europe. 
The  plant  is  well  suited  to  garden  use  and  does  not  intrude 
upon  other  plants.  It  does  not  do  well  in  the  dryest  places. 
The  cut  flowers  preserve  their  freshness  well.  Bot.  Mag.  266 
(poor).  Bot.  Cab.  1593.  Journ.  Hort.  Ill,  35,  p.  345.  Gard. 
Chron.  1883,  19:  788. 

76.  R.  Lambertianus  D.  DIETR.  Syn.  PI.  3:  316.     1843. 

Plant  swimming  :  leaves  lanceolate,  entire  or  subdenticulate, 
their  long  petioles  sheathing  the  stem  at  their  base :  flowers 
small,  yellow,  axillary  or  terminal ;  petals  obtuse,  longer  than 
the  stamens  and  sepals.  Wet  places.  Mexico.  R.  natans 
NEES,  ex  G.  Don,  Gen.  Syst.  i:  31,  1831  (not  C.  A. 
Meyer),  is  probably  a  form  of  this  with  leaves  sometimes  bifid. 
Mexico. 


494  MINNESOTA    BOTANICAL    STUDIES. 

77.  R.  arnoglossus  GREENE,  Pitt.  4:   143.     1900. 

Plant  tufted,  about  6  inches  high :  leaves  feather-veined, 
elliptic  and  elliptic-lanceolate,  entire ;  petioles  of  all  shorter 
than  the  blade,  sheathing  at  base  :  flowers  many,  large ;  petals 
5,  obovate,  obtuse,  commonly  persistent  with  the  sepals  :  akenes 
many  :  head  dense,  globose.  Subalpine  in  the  Ruby  mountains 
of  eastern  Nevada. 

78.  R.  unguiculatus  GREENE,  Pitt.  4:  142.     1900. 

Stem  i  foot  or  more  high,  solitary :  radical  leaves  I  or  2 
only,  erect,  elliptical,  or  obovate-elliptic,  acute,  entire  or  ob- 
scurely denticulate,  2  to  3  inches  long ;  petioles  as  long ;  stem 
leaves  narrower,  short-petioled  :  flowers  2  to  4,  or  more  in  the 
large  plants  ;  peduncles  long,  puberulent,  naked ;  petals  about 
10,  persistent,  narrow,  claw  y2  line  long ;  sepals  narrow, 
spreading,  deciduous :  akenes  glabrous,  obliquely  obovoid, 
slightly  compressed ;  beak  stout,  slightly  recurved  :  head  de- 
pressed globose.  11,500  feet.  Southern  Colorado.  C.  F. 
Baker,  August  28,  1899. 

79.  R.  ambigens  WATS.  Bibl.  Index,  i :   16.     1878. 

R.  Flammula  PURSH,  Fl.  2:  391.      1814.     Not  Linn. 

R.  Lingua  PURSH,  1.  c.     Not  Linn. 

R.  alismafolius  BENTH.  PI.  Hartw.  295.      1848.     In  part. 

Not  Geyer. 
R.  obtusiusculus  BRITTON,  111.   Fl.   2:    76.     1895.      Not 

Raf. 

Plant  2  to  3  feet  high,  stout,  glabrous  or  nearly  so,  erect,  but 
sometimes  rooting  at  the  lower  nodes,  hollow :  leaves  usually 
on  short  petioles  with  broad,  membranous,  sheathing  bases ; 
blades  lanceolate  with  tapering  bases,  serrate,  denticulate  or 
entire,  2  to  4  inches  long:  petals  5  to  7,  yellow,  2  to  3  lines 
long  ;  sepals  shorter  :  akenes  small,  obliquely  oval,  compressed, 
thickened  along  one  margin ;  beaks  subulate,  narrow,  erect  or 
little  curved,  nearly  as  long  as  akene  body :  head  of  fruit  glo- 
bose. Wet  grass  lands.  Mountains  of  Georgia  and  Tennes- 
see to  Missouri,  north  and  east  to  Canada  and  New  England. 

Var.  obtusiusculus  n.  var. 
R.  obtusiusculus  RAF.  in  Desv.  Journ.  Bot.  1 :   225.     1808. 

Differs  from  the  type  in  its  slender,  straight,  erect  stem  :  its 
single  root,  like  an  annual,  and  its  linear-lanceolate  sepals. 


Davis:    RANUNCULI  OF  NORTH  AMERICA.  495 

So.   R.  Madrensis  ROSE,  Cont.  Nat.  Herb.  5:   199.     1899. 

Plant  erect,  rather  slender,  6  to  12  inches  high,  glabrous  on 
lower  parts ;  i-4-flowered ;  radical  leaves  i  to  2  inches  long, 
petioled,  linear  to  linear-oblong,  with  coarse  distant  teeth,  ob- 
tuse ;  base  cuneate  ;  stem  leaves  reduced  to  bracts,  simple  or 
3-lobed  :  flowers  yellow,  on  long  slender  peduncles  which  are 
hairy  near  the  flower  ;  receptacle  hairy  ;  petals  about  10,  obovate 
to  oblong,  3  to  4  lines  long  :  akenes  hardly  i  line  long,  com- 
pressed, glabrous  ;  beak  as  long,  slender.  Sierra  Madre  moun- 
tains, between  Santa  Gertrudis  and  Santa  Teresa,  Tepic  Ty., 
and  in  Zacatecas.  Altitude  7,400  to  10,000  feet. 

81.  R.   alismaefolius    GEYER,    in    Benth.    PI.    Hartw.    295. 

1848. 

JR.  Flammula  HOOK.  Lond.Journ.  Bot.  6:  66.     1847. 
R.  Bolanderi  GREENE,  Bull.  Calif.   Acad.   Sci.    2  :  58. 

1886. 
R.  Hartwegi  GREENE,  Erythea,  3:  45.     1895. 

Roots  fibrous,  fascicled  :  plant  erect,  usually  robust,  6  to 
15  inches  high,  branching  or  nearly  simple,  slightly  pubescent 
on  peduncles  :  leaves  oblong  to  lanceolate,  tapering  at  base, 
entire  or  denticulate,  i  to  3  inches  long ;  petioles  short  and 
broad,  sheathing  at  base ;  upper  stem  leaves  sessile :  petals 
yellow,  6  to  10,  obovate,  4  to  6  lines  long  ;  sepals  much  shorter, 
reflexed :  akenes  compressed,  smooth ;  beak  short,  often 
hooked  :  head  of  fruit  nearly  globose.  Wet  grounds.  British 
Columbia  and  Colorado  to  California. 

Var.  Calthaeflorus  n.  var. 
R.    Calthceflorus  GREENE,  Erythea,  3:  45.      1895. 

Leaves  repand-denticulate,  and  much  broader  than  the  type : 
flowers  and  petals  not  different.  Colorado,  in  boggy  ground, 
8,000  feet. 

82.  R.  alismellus  GREENE,  Fl.  Francis,  2:  97.     1892. 

R.  alismafolius  var.  alismellus  GRAY,  Proc.  Am.  Acad. 

7:   327.      1868. 
R.  altsmafolius  var.  montanus  WATS.  Bot.  King  Exp.  7. 

1871. 

Much  like  R.  alism<efolius ,  but  usually  very  slender,  dwarf, 
often  nearly  scapiform :  leaves  lanceolate-elliptical  to  ovate  : 
petals  smaller,  3  lines  long.  Plants  often  form  a  thick  cover- 


496  MINNESOTA    BOTANICAL    STUDIES. 

ing  over  the  wet  ground.      High   altitudes,  Sierra   and    Trinity 
Mts.,  California  to  Colorado  and  northward. 
Var.  Populago  n.  var. 

R.  Populago  GREENE,  Erythea,  3  :   19.     January,  1895. 
R.    Cusickii  JONES,  Proc.  Calif.  Acad.  Sci.,  Ser.  2,  5  :  615. 

October;   1895. 

Like  the  type,  but  with  radical  leaves  ovate,  cordate ;  mar- 
gins slightly  wavy.  Southwestern  Oregon,  Idaho. 

83.  R.  Lemmoni  GRAY,  Proc.  Am.  Acad.  10  :  68.     1874. 

Stems  scapose,  tufted,  5  to  10  inches  high  :  plant  villous- 
pubescent  on  lower  parts  :  leaves  rather  thick,  lanceolate,  en- 
tire :  flowers  i  or  2  on  a  stem  ;  petals  about  three  lines  long, 
obovate  to  oblong  :  akenes  turgid,  villous-pubescent,  borne  in 
an  oval  head.  Rare.  Truckee  and  east  part  of  Sierras,  Cali- 
fornia. 

84.  R.  oblongifolius  ELL.  Sketch  2  :  58.     1824. 

R.  Flamniula  MICHX.  Fl.  i  :   221.      1803.     Not  Linn. 
R.  Flammula  var.  laxicanlis  TORR.   &  GRAY,  Fl.    i  :   16, 

1838. 

R.  pusillus  var.  oblongifolius  TORR.  &  GRAY,  1.  c.  17. 
R.  laxicanlis  DARBY,  Bot.  S.  St.  204.      1855. 

Annual ;  about  i  to  2  feet  high,  erect  or  ascending,  rarely 
rooting  at  the  lower  nodes,  branched  above,  many-flowered : 
leaves  shaped  nearly  as  in  R.  -pusillus^  or  sometimes  broader : 
petals  5,  yellow,  longer  than  the  sepals  ;  stamens  many  :  akenes 
few,  often  globular  or  slightly  flattened,  smooth  or  minutely 
punctate  ;  style  deciduous  :  head  of  fruit  globose.  April  to  Sep- 
tember. Wet  grounds.  Florida  to  southern  Virginia,  west  to 
southern  Missouri  and  Texas. 

85.  R.    hydrocharoides    GRAY,   Mem.  Am.    Acad.    5:  306. 

1855- 

Stems  ascending,  5  to  10  inches  high,  rooting  at  the  lower 
nodes,  with  creeping,  fistulous  branches  at  the  base :  leaves 
mostly  long-petioled,  entirely  or  nearly  so,  usually  less  than  i 
inch  long,  rather  succulent ;  basal  leaves  round-cordate  to  oval, 
blending  into  the  form  of  the  upper  ones  which  are  obovate  to 
spatulate :  petals  5  or  more,  2  to  3  lines  long ;  sepals  much 
shorter  :  akenes  small ;  beak  narrow,  short :  head  of  fruit  small, 
globose.  In  standing  water  and  wet  soil.  Southwestern  Ari- 


Da-vis:   RANUNCULI  OF  NORTH  AMERICA.  497 

jona  into  adjacent  California,  and  in  Lower  California,  at  La 
^huparosa  and  Sierra  de  la  Laguna.  Flowers  appearing  very 
ate  at  high  altitudes. 

86.  R.  samolifolius  GREENE,  Pitt.  3:  13.     1896. 

Much  like  R.  hydrocharoidcs.  Leaves  entire,  obtuse,  ob- 
anceolate,  petioled ;  upper  ones  oval  or  obovoid :  petals  ob- 
>vate  ;  sepals  round-ovoid,  spreading  :  akenes  like  that  species. 
Sigh  altitude,  Mt.  Shasta,  southward. 

87.  R.  stolonifer  HEMSL.  Diag.  PI.  Nov.  17.     1879. 

Plant  small,  entirely  glabrous,  spreading  by  stolons  :  stem 
srect,  2  to  6  inches  or  less  :  leaves  subentire  or  sometimes  cre- 
late,  the  radical  ones  long-petioled,  reniform  or  roundish  ellip- 
ical  to  lanceolate-oblong  ;  blade  3  to  12  lines  long,  petiole  i  to 
i,  with  base  membranous  and  dilated  ;  stem  leaves  sessile,  nar- 
•ow  :  flowers  small,  yellow,  long-peduncled  ;  receptacle  conical, 
glabrous  ;  sepals  oblong,  i  to  i  ^  lines  long  ;  petals  5  or  6,  ob- 
ong-elliptical,  about  i^/(  lines  long,  the  long  claw  with  a  con- 
spicuous nectary  ;  stamens  longer  than  petals,  filaments  dilated  : 
ikenes  much  compressed,  slightly  margined,  glabrous :  head  of 
rruit  globose.  Near  Morales  in  San  Luis  Potosi,  Mex.,  6,000 
o  8,000  feet  altitude. 

88.  R.  vagans  WATS.  Proc.  Am.  Acad.  26:   131.     1891. 

Plant  low,  glabrous,  spreading  by  elongated  stolons :  leaves 
larrowly  lanceolate,  or  the  lowest  ovate-lanceolate,  entire  or 
•vith  a  few  often  slender  teeth  toward  the  apex :  petals  8  to  10, 
Dblong-obovate,  about  2^  lines  long,  a  prominent  nectar  pore 
ibove  the  narrow  claw  ;  sepals  little  over  half  as  long  :  akenes 
smooth,  in  a  dense  globose  head  2^4  lines  in  diameter.  Flor 
ie  Maria,  State  of  Mexico.  Aug.,  1890.  Pringle  no.  3177. 

89.  R.  reptans  LINN.  Sp.  PI.  549.     1753. 

R.  filiformis  lA\cw&.  Fl.  i:  320.     1803. 
R.  reptans  var.  filiformis  DC.  Syst.  1 :   248.     1818. 
R,  Flammiila  var.  filiformis  HOOK.  Fl.  I  :   n.      1829. 
R.  Flammulavar.  reptans  E.  MEYER,  PL  Labr.  96.    1830. 

Stem  prostrate,  rooting  at  the  nodes,  pubescent  or  nearly 
glabrous  :  leaves  linear-lanceolate  to  spatulate,  usually  entire, 
[  to  2  inches  long,  narrowed  into  the  petiole  :  peduncles  ascend- 
ng,  i  to  3  inches,  each  terminated  by  a  single  flower ;  petals 


498  MINNESOTA    BOTANICAL    STUDIES. 

4  to  7,  bright  yellow,  2  to  3  lines  long,  exceeding  the  sepals; 
stamens  many  :  akenes  flattened  somewhat ;  beak  minute,  sharp. 
Coast  of  Arctic  America,  Newfoundland  ;  near  ponds  and  lakes, 
New  Jersey  to  California  :  Greenland,  Europe,  Asia. 

Var.  Gormani  n.  var. 
R.  Gormani  GREENE,  Pitt.  3:  91.      1896. 

Like  the  type,  but  with  leaves  broadly  ovate  or  deltoid-ovate, 
acute,  few-toothed,  6  to  8  lines  long.  Near  Crater  lake,  south- 
ern Oregon. 

90.  R.   Unalaschensis   BESS,   ex   Ledeb.   Fl.   Ross,   i :  32. 

1841. 

R.  Flammula  var.  intermedius  HOOK.  Fl.  I  :   n.      1829. 
R.  reptans  var.  intermedius  TORR.  &  GRAY,   Fl.  I  :   16. 

1838. 
R.  Flammula  var.   Unalaschensis  LEDEB.  ex  Regel,  Bull. 

Soc.  Nat.  Mosc.  34:  pt.  2  :  41.     1861. 
R.  reptans  var.  strigulosus  FREYN,  Deutsch.  Bot.  Monats. 

8:   181.      1891. 
R.    intermedius    HELLER,     Bull.    Torr.    Club,   25 :    280. 

1898.     Not  Poir.  nor  Eaton. 

This  differs  from  R.  reptans  in  its  more  robust  habit,  longer 
peduncles,  leaves  larger,  sometimes  being  3  to  5  inches  long 
and  2  to  6  lines  wide.  Newfoundland  past  the  Great  Lakes  to 
Oregon  and  California,  northward.  Europe.  Asia. 

91.  R.  microlonchus  GREENE,  Erythea,  4:  122.     1896. 

Allied  to  R.  reptans,  often  more  hairy ;  stem  slender,  some- 
what ascending,  i -few-flowered  :  radical  leaves  in  a  tuft,  shaped 
as  in  that  species  or  a  little  broader ;  stem  leaves  few,  short 
petioled  to  subsessile  :  flowers  4  lines  broad ;  petals  5  to  8,  ob- 
tuse ;  sepals  spreading  :  akenes  as  in  that  species  but  with  a 
short,  stout,  blunt  beak :  head  of  fruit  depressed-globose. 
Northern  Idaho. 

92.  R.  trachyspermus  ENGELM.  ex  Gray  PL   Lindh.  i :  3. 

1850. 

R.  trachyspermus  var.  angustifolius  ENGELM.  1.  c. 

Annual :  plant  glabrous,  y2  to  2  feet  high,  sometimes  rooting 

at  the  lower  nodes  :  leaves  slender-petioled,    oblong  to  linear 

lanceolate,  entire  or  denticulate,  bases  often  tapering  ;  petioles 

of  stem  leaves  expanded  near  the  bases  :  peduncles  rather  short ; 


Davis:    RANUNCULI  OF  NORTH  AMERICA.  499 

petals  i  to  3  or  5,  pale  yellow,  about  i  line  long  ;  stamens  only 
5  to  10 :  akenes  oblong,  hardly  compressed,  slightly  margined ; 
beak  very  short :  head  of  fruit  oblong.  Low,  wet  places,  Dal- 
las, Tex.,  south  and  southeast. 

93.  R.  pusillus  POIR.  in  Lam.  Encycl.  6:  99.     1804. 
R.  Flammula  WALT.  Car.  159.     1788.     Not  Linn. 
R.  Bonariensis  POIR.  in  Lam.  Encycl.  6:  102.     1804. 
R.  humilis  PERS.  Syn.  2:  102.      1807. 

Annual :  plant.  */2   to  2  feet  high,  slender,  weak,  branching, 
glabrous  :  basal  and  lower  stem-leaves  mostly  ovate,  petiolate ; 
others  nearly  sessile,  linear  or  lanceolate  ;  all  entire  or  minutely 
toothed  :   petals  yellow,  few,  barely  exceeding  the  sepals ;  sta- 
mens i  to  10  :  akenes  obovate,  tipped  with  the  base  of  the  style  : 
head    of    fruit    globose.     Marshy    ground,    New   York,    New 
Jersey,  to  Florida,  through  Gulf  region  to  Texas  and  Missouri. 
Var.  Lindheimeri  GRAY,  Proc.  Am.  Acad.  21 :  367.     1886. 
R.  trachyspcrmus  var.  Lindheimeri  ENGELM.  ex  Gray  PI. 

Lindh.  i  :  3.      1850. 

R.  Biolettii  GREENE,  Pitt.  2:  225.     1892. 
Low,  rarely  a  foot  high :  akenes  more  papillose-roughened 
than  in   the   type.     Middle  coast  of  California  to   Galveston, 
Tex.,  and  New  Orleans. 

94.  R.  Anderson!  GRAY,  Proc.  Am.  Acad.  7:  327.     1868. 
Oxygraphis  Andersoni  FREYN,  Flora,  70:  140.     1887. 

About  6  to  8  inches  high,  stem  one-leaved  or  a  naked  scape  : 
basal  leaves  rather  thick,  2  to  3  times  ternately  or  pedately  di- 
vided or  parted,  lobes  linear  to  lanceolate :  flowers  i  or  2 ; 
sepals  glabrous ;  petals  ^  inch  long,  pink  or  rose,  orbicular, 
obovate  or  flabellate,  claws  narrow :  akenes  compressed,  but 
wholly  utricular  with  membranous  walls,  oblique  obovate  to  or- 
bicular, y^  inch  long,  a  very  narrow  membranous  margin  ;  apex 
abruptly  sharpened  with  a  very  short  style.  Boise  City,  Idaho, 
Salt  lake,  Utah,  to  eastern  Sierras  of  California  and  Nevada. 

95.  R.  juniperinus  JONES,  Proc.  Cal.  Acad.  Sci.,  Ser.  2,  5: 

616.     1895. 
R.  Andersoni  var.  tenettusWATS.  Bot.  King  Exp.  7.  /.  /, 

f.  8-10.     1871. 

Plant  taller  and  more  slender  than  R.  Andersoni;  usually 
branched  once,  i-leaved  and  2-flowered :  radical  leaves  more 


500  MINNESOTA    BOTANICAL    STUDIES. 

finely  dissected  than  in  that  species  :  petals  white  or  rose-purple 
without:  akenes  flat,  not  inflated,  very  small,  only  i  to  il/2  lines 
long.  Rocky  places  in  woods  of  Utah. 

96.  R.  glacialis  LINN.  Sp.  PI.  553.     1753. 

Roots  fibrous  :  plant  low :  lower  leaves  petioled,  others  ses- 
sile and  involucrate,  all  3-parted  or  trifid  and  again  lobed ;  up- 
per ones  often  villous  :  flowers  i  to  3,  white  or  reddish;  petals 
obovate  to  cuneate,  blunt ;  sepals  shorter,  very  densely  hairy. 
Summer.  Mountains  of  Europe,  Arctic  regions,  Greenland. 
Garden  45,  p.  28  ;  48  p.  501. 

KUMLIENIA  GREENE,  Bull.  Calif.  Acad.  i :  337.     1886. 
(Named  for  the  late  botanist,  Prof.  Kumlien  of  Wisconsin.) 

Low  perennials ;  stem  nearly  leafless,  i-2-flowered  :  leaves 
mostly  radical,  rounded  and  lobed:  sepals  5  to  7,  white,  con- 
spicuous; petals  5,  small,  oval,  fleshy,  with  nectariferous  pit 
and  slender  claw ;  stamens  and  pistils  many  :  akenes  lanceolate, 
acuminate,  compressed,  membranaceous,  and  utricular,  ob- 
scurely i-nerved  on  the  sides;  style  hooked,  persistent;  seed 
much  shorter  than  the  cell.  A  monotypic  genus  of  narrow  dis- 
tribution. Of  it  Greene  remarks  that  it  has  the  general  aspect 
of  Ranunculus ;  flowers  of  Caltha,  with  nectary-like  petals  of 
Helleborus,  the  utricular  fruit  peculiar.  This  is  section  PSEUDA- 
PHANTOSTEMMA  of  Gray,  under  Ranunculus. 

K.  hlstricula  GREENE,  1.  c. 

Ranunculus  hystriculus  GRAY,  Proc.  Am.  Acad.  7:  328. 
1868. 

Stems  4  to  10  inches  high,  bearing  i  or  2  3-lobed  leaves : 
radical  leaves  round-reniform,  with  5  broad  rounded  lobes ; 
petioles  long  :  flowers  i  or  2  :  akenes  3  lines  long  including  the 
style.  April  to  June.  Portland,  Ore.,  east  to  the  Sierra  Neva- 
das.  Rare. 

FICARIA  HUBS.  Fl.  Angl.  213.     1762. 
(Latin  for  fig,  referring  to  the  thickened  roots.) 

Perennial  herbs  with  tuberous  roots ;  plants  all  glabrous ; 
stems  branched  or  simple  :  leaves  petioled,  entire  or  toothed,  cor- 
date :  flowers  large,  solitary,  either  axillary  or  terminal ;  sepals 
3  to  5,  deciduous;  petals  about  8  (7  to  12),  yellow,  or  red  at 


Davis:    RANUNCULI  OF  NORTH  AMERICA.  501 

the  base  ;  carpels  numerous,  blunt,  not  wrinkled  nor  ribbed, 
cotyledon  only  i  :  akenes  borne  in  a  head.  A  genus  of  only 
about  three  species  natives  of  Europe  and  Asia.  The  follow- 
ing one  is  naturalized  here  : 

F.  ranunculoides  MOENCH,  Meth.  215.     1794. 

Ranunculus  Ficaria  LINN.  Sp.  PI.  550.     1753. 

F.  verna  HUDS.  Fl.  Angl.  i  ed,  214.      1762. 

F.  -polypetala  GILID.  Fl.  Lituan.  2:   259.      1781. 

F.  Ficaria  KARST.  Deutsch.  Fl.  565.  1880-83.  (Not  bi- 
nomial.) 

F.  communis  DuM.-CouRS.  Bot.  Cult.  2  ed  .4 :  445.     1811. 

F.  calt/HEfoItaR.vicHB.  Fl.  Germ.  Excurs.  718.      1830-32. 

F.  grandiflora  ROBERT,  Cat.  Toulon,  57:   112.      1838. 

F.  Robcrti  F.  SCHULTZ,  Arch.  Fl.  346.     1848. 

F.  ambigua  BOR.  Fl.  Cent.  Fr.  3  ed.  2  :  20.      1857. 

F.  nudicaulis  KERN,  in  Oestr.  Bot.  Zeitschr.  13 :  188. 
1863. 

F.  intermedia  SCHUR.  Enum.  PL  Transs.  14.      1866. 

F.  transsilvanica  SCHUR.  1.  c.  14. 

F.  aperata  SCHUR.  inVerh.  Naturf .  Ver.  Bruenn.  15:  231. 
1877. 

F.  Holubyi  SCHUR.  1.  c.  32. 

F.  rotundifolia  SCHUR.  1.  c.  32. 

Stem  scape-like,  or  i-2-leaved,  about  5  inches  high  :  leaves 
ovate  cordate,  obtuse,  crenate,  i  to  2  inches  long ;  petioles 
broad  :  flowers  yellow  ;  petals  8  or  9  ;  sepals  3  :  head  of  fruit 
globose  :  akenes  beakless,  truncate.  Run  wild  on  Long  is- 
land;  Staten  island;  Hingham,  Mass.;  Richmond  Co.,  N. 
Y.,  and  near  Philadelphia.  Regne  Vegetale  9:  6. 

CYRTORHYNCHA  NUTT.  ex  Torr.  &  Gray,  Fl.  i :  26.     1838. 
(From  the  Greek,  meaning  curved-beak.) 

Slender,  erect,  perennial  herbs,  with  fibrous  roots :  leaves 
lobed:  flowers  small,  yellow;  sepals  5,  spreading  deciduous; 
petals  5,  narrowly  spatulate  or  oblong,  pit  at  base,  small; 
stamens  many :  akenes  terete,  longitudinally  ribbed ;  style  in- 
curved. A  monotypic  genus.  Section  CYRTORHYNCHA  Gray, 
under  Ranunculus. 


502  MINNESOTA    BOTANICAL    STUDIES. 

C.  ranunculina  NUTT.  1.  c. 

Ranunculus  Nuttallii  GRAY,  Proc.  Acad.  Phil.  56.      1863. 
R.  ranunculinus  RYDBERG,  Bot.  Surv.  Neb.  3  :  23.     1894. 

Glabrous,  6  to  10  inches  high  :  leaves  usually  narrowly  lobed  ; 
basal  ones  long-petioled  ;  stem  leaves  few,  beneath  the  branches  : 
flowers  several,  somewhat  corymbose  :  akenes  tipped  with  the 
persistent,  slender,  recurved  style.  Spring  and  Summer.  Ne- 
braska to  Wyoming.  C.  neglecta^  in  Greene's  herbarium,  is  a 
form  with  roots  more  succulent ;  stems  and  leaves  like  the 
above  ;  petals  perhaps  a  little  larger. 

ARCTERANTHIS  GREENE,  Pittonia,  3:   190.     1897. 

(Combination  of  Arctic  and  Eranthis:  in  allusion  to  its  habitat 
and  resemblance  to  Eranthis •,  or  Cammarum). 

A  monotypic  genus  of  perennial  herbs  :  roots  rather  fascicled 
or  clustered  on  a  short  caudex :  leaves  mostly  radical,  rounded 
and  lobed :  flowers  solitary ;  sepals  5  ;  petals  10 ;  stamens 
many :  akenes  in  a  head,  longitudinally  ribbed,  beaks  reflexed. 
Part  of  Section  CYRTORHYNCHA  Gray,  Syn.  Fl.  i  :  23,  under 
RANUNCULUS. 

A.  Cooleyae  GREENE,  1.  c.  190.  pi.  j. 

Ranunculus    Cooleyce.  VASEY  &  ROSE,  Contr.  Natl.  Herb. 

I  :   289,  pi.  22.      1893. 
Kumlienia  Cooleyce  GREENE,  Erythea,  2:   193.     1894. 

Plant  glabrous,  3  to  8  inches  high :  scape  i-2-flowered, 
sometimes  bearing  a  small  leaf  near  the  middle,  extending 
above  the  leaves  when  in  fruit ;  basal  leaves  many,  orbicular, 
one  inch  or  more  across,  deeply  3-parted,  and  again  lobed  and 
toothed,  glandular  tips  to  the  teeth ;  petioles  broadened  or 
sheathing  at  the  base  :  flowers  yellow  ;  sepals  oblong,  obtuse, 
deciduous ;  petals  oblong,  obtuse,  tapering  into  a  slender  claw 
at  base :  carpels  compressed  laterally,  i-3-nerved  on  each 
side ;  reflexed  style  short ;  ovule  erect.  August.  Mountain 
tops  near  Juneau,  and  St.  Elias  Alps,  Alaska. 

OXYGRAPHIS  BUNGE  Verz.  Suppl.  Fl.  Alt.  46.     1836. 
(From  Greek,  meaning  sharp-style.) 

Trailing  and  running  perennial  herbs,  with  fibrous  roots : 
leaves  crenate-dentate  or  lobed,  long-petioled,  glabrous  :  flow- 


Davis:   RANUNCULI  OF  NORTH  AMERICA.  503 

ers  small,  yellow,  one  to  several  on  scape-like  stems ;  sepals 
often  5,  deciduous;  petals  5-12,  pit  at  base  small;  stamens 
many :  akenes  compressed,  somewhat  swollen,  thin-walled, 
striate  with  simple  or  branched  nerves  :  head  of  fruit  oblong  or 
oval.  Several  species,  but  the  following  only  is  found  in 
America. — Section  HALODES,  Gray,  Proc.  Am.  Acad.  21 : 
366,  under  Ranunculus. 

0.  Cymbalaria  PRANTL,  in  Engl.  &  Prantl,  Nat.  Pfl.  Fam. 

3:  Abt.  2,  63.     1891. 

^Ranunculus  salnginosus  PALL.  Reise.  3  :   263.      1776. 
R.  Cymbalaria  PURSH,  Fl.  2:  392.     1814. 
R.  halophilus  SCHLECHT.  Animad.   Ranunc.  i:    23.  t.  4. 

/.  /.     i8iQ. 

R.  tridentatus  H.B.K.  Nov.  Gen.  &  Sp.  PL  5  :  42.     1821. 
R.  Cymbalaria  var.  alpmus  HOOK.  Fl.  i:    n.     1833. 
Cyrtorhyncha    Cymbalaria  BRITTON,   Mem.  Torr.   Club, 

5:   161.      1894. 

Leaves  orbicular  to  ovate,  cordate  or  truncate  at  base,  one 
inch  long  or  even  much  less  :  scapes  short ;  receptacle  elongat- 
ing in  fruit :  akenes  minutely  sharp-pointed.  Summer.  Shady 
shores  and  moist  saline  or  salt  grounds.  Arctic  sea  coasts  to 
New  Jersey,  west  to  Minnesota,  thence  south  and  west.  Also 
Mexico,  South  America,  Greenland,  Asia. 

INDEX  TO  NAMES  OF  RANUNCULUS. 
(References  to  other  genera  are  indicated  by  their  initials.) 

aborti-vus  Hook.,  35.  affinis  Torr.,  49. 

abortivus  Linn.,  32.  affinis  var.  cardiophyllus,  42. 

abortivus  var.  australis,  32.  affinis  var.  lasiocarpus,  42. 

abortivus  var.  encyclus,  32.  affinis  \a.r.'leiocarpus,  42. 

abortivus  var.  grandiflorus,  32.      ajfinis  var.  micropetalus,  42. 

abortivus  var.  Harveyi,  32.  affinis  var.  validus,  42. 

abortivus  var.  micranthus,  10.  alceus,  6. 

aconitifolius,  73-  alisnuzfolius  Benth.,  79- 

acris  Hook.,  29.  alismcEfolius  Geyer,  81. 

acris  Linn.,  40.  alismcefolius  var.  alismellus,  82. 

acris  var.  Deppii,  30.  alismcefolius    var.     calthceflorus, 

acriformis,  29.  Si. 

adoneus,  58.  alismcefolius  var.  montanus,  82. 

ajfinis  R.  Br.,  42.  alismellus,  82. 


504 


MINNESOTA    BOTANICAL    STUDIES. 


alismellus  var.  Populago,  82. 

Allegheniensis,  35. 

alpeophilus,  45. 

amarittoi  25. 

ambigens,  79. 

ambigens  var.  obttisiusculus,  79. 

amcenus  Gray,  58. 

amoenus  Ledeb.,  42. 

amplexicaulis,  75. 

Andersoni,  94. 

Andersoni  var.  tenellus,  95. 

apricus,  1 1 . 

aquatilis,  =  B.  2. 

aquatilis  /?,  =  B.  i. 

aquatilis  var.  br  achy  pus,  =  B.  3. 

aquatilis  var.  ccespitosus,  =  B.  3. 

aquatilis  var.  confervoides,  =  B. 

3- 
aquatilis  var.    divaricatus,  =  B. 

i. 

aquatilis  var.  flaccidus,  =  B.  3. 
aquatilis    var.    heterophyllus,  = 

B.  2. 

aqziatilis\2it.  hispidulus,  =  B.  2. 
aquatilis  var.  Lobbii,  =  B.  5. 
aquatilis  var.   longirostris,  =  B. 

i. 

aquatilis  var.  submersus,  =  B.  3. 
aquatilis  var.    stagnatalis,  =  B. 

i. 
aquatilis    var.    trichophyllus,  = 

B.3. 

arcticus,  42. 
arcuatus,  37. 
Arizonicus  Greene,  49. 
Arizonicus  Lemmon,  49. 
Arizonicus  var.  subajfinis  Gray, 

49- 
Arizonicus  var.  subajfinis  Greene, 

42. 
Arizonicus  var.  subsagittatus, 

49- 
arnoglossus,  77. 


arvensis,  I . 

Ascherbornianus,  28. 

Asiaticus,  31. 

auricomus,  42. 

auricomus  var.   Cassubicus,  48. 

Austince,  64. 

Beckii,  70. 

Belvisii,  12. 

Biolettii,  93. 

Bloomer  i,  15. 

Bolanderi,  81. 

Bonariensis,  93. 

Bongardi,  37. 

Bongardt  var.  Douglasii,  37. 

Bongardi  var.  tenellus^  37. 

brevicaulis,  48,  64. 

bulbostis,  17- 

Canadensis,  7. 

canus,  24. 

canus  var.  Blankinshipii,  24. 

canus  var.  hesperoxys,  24. 

Calif ornicus,  30. 

Calif ornicus  var.  canus,  24. 

Calif ornicus  var.  crassifolius^o, 

Californicus  var.  latilobus,  30. 

Calif  ornicus  var.  Ludovicianus, 

3°. 

calthceflorus,  Si. 
cardiophyllus,  42. 
cardiophyllus  var.  pinctor um,  42. 
carpaticus,  50. 
confervoides,  =  B.  3. 

Cooleyce,  =  A. 
corthuscefolius,  5 1 . 

Chilensis,  15. 
ciliosus,  38. 
circinatus,  =  B.  i. 

Clintonii,  26. 

Cusickii,  82. 

Cymbalaria,  =  O. 

Cymbalaria  var.   alpinus,    =  O. 
delphinifohus,  H.B.K.,  23. 
delphinifolius  Torr.,  70. 


Davis:   RANUNCULI  OF  NORTH  AMERICA. 


505 


delphinifolitis  T.  &  G.,  30. 
dichotomus,  20. 
digitatus,  63. 
dissect  us,  39. 
divaricatus,  =  B.  i. 
Donianus,  53. 
Douglasii,  37. 
Drummondi,  60. 
Ear  lei,  37. 
Eiseni,  38. 
ellipticus,  64. 
ercmogenes,  34. 
cremogenes  var.  degener,  34. 
Eschscholtzii,  44. 
eximius,  45. 
fascicularis  Britton,  16. 
fascicularis  Muhl.,  1 1 . 
fascicularis  Schlecht.,    12. 
fascicularis  Wats.,  7- 
fascicularis  VM:.  Deforests,  n. 
Ficaria,  =  F. 
filiform  is,  89. 
flaccidtis,  =  B.  3. 
Flammula  Hook.,  Si. 
Flammula  Michx.,  84. 
Flammula  Pursh,  79. 
Flammula  Walt.,  93. 
Flammula  var.  filiformis,  89. 
Flammula  var.   intermedius,  90. 
Flammula  var.  laxicaulis,  84. 
Flammula  var.  reptans,  89. 
Flammula  var.  Unalaschensis^o. 
flaviatilis,  7°- 

Galeottii,  5. 
geoides,  55. 
glaberrimus,  64. 
glaberrimus  var.  ellipticus,  64. 
glacialis,  96.  f 

Gmelini,  7 r  • 

Gormani,  89. 

Grayanus,  =  B.  2. 

Grayi,  60. 

Greenei,  37. 


halophilus,  =  O. 
Hartivegi,  8 1 . 
Harvey  i,  32. 
hebecarpus,  4. 
hebecarpus  var.  pusilhts,  4. 
hederaceus  Linn.,  =  B.  4. 
hederaceus  var.  =  B.  5. 
hederaceus  var.   Lobbii,  =  B.  5. 
hesperoxys,  24. 
hirsutus,  9. 
hispidus  Hook.,  8. 
hispidus  Michx.,  16. 
hispidus  Pursh,    7- 
hispidus  var.  Oreganus,  8. 
Hooker  i  Regel,  60. 
Hookeri  Schlecht.,  13. 
Ho-wellii,  38. 
humilis  D.  Don.,  53. 
humilis  Pers.,  93. 
hydrocharis  sub.  sp.  Lobbii,   = 
B.  5. 

hydrocharoides,  85. 

hyperboreus,  68. 
hyperboreus  var.  natans,  67. 

hystriculus,  =  K. 

Icelandicus,  18. 

inamoenus,  42. 

intermedius,  90. 
juniperinus,  95. 

lacustris,  7°- 

Lambertianus,  j6. 

L  a  ngsdo  rfii,  7 x  • 

lanuginosus,  36. 

Lapponicus  Linn.,  69. 

Lapponiciis  Oed.,  61. 

laxicaulis,  84. 

Lcmmoni,  83. 

limosus,  71. 

Lingua,  79. 

Llavanus,   2 1 . 

Lobbii,  =  B.  5. 

longipedunculatus,  52. 

longirostris,  =  B.  i. 


506 


MINNESOTA    BOTANICAL    STUDIES. 


lucidus,  12. 
Ludovicianus,  30. 
Lyatti,  37. 
Macauleyi,  65. 

Macounii,  8. 

Macounii  var.  Oreganus,  8. 

macranthtis  Scheele,  22. 

macranthus  Wats.,  19. 

Madrensis,  80. 

Marilandicus,  16. 

maximus,  19. 

McCattai,  41. 

Mexicanus,  55. 

micranthus,  10. 

microlonchtis,  91. 

Missouriensis,  72. 

Montanensis,  29. 

montanus,  50. 

montanus  var.  dentatus,  50. 

mtilticaulis,  54. 

multifidus,  70. 

?nultifidus  var.  limosus,  71- 

multifidus  var.  repens,  71. 

mtiltifidus  var.  terrestris,  70. 

muricatus,  2. 

natans  C.  A.  Meyer,  67. 

natans  Nees,  76. 

Nelsoni,  38. 

Nelsoni  var.  glabriusculus,    37. 

Nelsoni  var.  tenellus,  37- 

nitidus,  32. 

nivalis  Linn.,  66. 

nivalis  Rep.  not  Linn.,  65. 

nivalis  var.  Eschscholtzii,  44. 

Nuttallii,  =  C. 

oblongifolins,  84. 

obtusiusculus  Britton,  79- 

obtusiuscuhis  Raf.,  79- 

occidentalis  Gray,  38. 

occidentalis  Nutt.,  38. 

occidentalis  var.  brevistylus,  38. 

occidentalis  var.  Eiseni,  37,  38. 

occidentalis  var.  Hovoellii,  38. 


occidentalis  var.  Lyalli,  37. 
occidentalis  var.  parviflorus,  37. 
occidentalis  var.  Rattani,  38. 
occidentalis  var.  robusttis,  38. 
occidentalis  var.  tenellus,  37. 
occidentalis  var.   ultramontamis, 

33- 

ocreatus,  44. 
Oreganus,  8. 
ornithorhynchus,  19. 
orthorhynchus,  19. 
orthorhynchus  var.  alpinus,  58. 
orthorhynchus  var.  platyphyllus, 

19. 

ovalis,  48. 
oxynotus,  62. 
Pallasii,  74. 
palmatus,  27. 
pantothrix,  =  B.  3. 
parviflorus  Linn.,  3. 
parvijlorus  var.  T.  &  G.,  4. 
parmilus,  9. 
pedatifidus  Hook.,  60. 
pedatifidus  J.  E.  Smith,  42. 
pedatifidus    var.     cardiophyllus, 

42. 

pedatifidus  var.  pinetorum,  42. 
Pennsylvanicus,  J. 
petiolaris,  57. 
Phitonotus  Ehrh.,  9. 
Philonotus,  Pursh,  12. 
pilosus,  14. 
Populago,  82. 
Porteri,  =  B.  3. 
prostratus,  26. 
Purshii  Richards,  71. 
Purshii  To  IT.,  67. 
Piirshii  var.  aquatilis,  7°- 
pusillus  Ledeb.,  71. 
pusillus  Poir.,  93. 
pusillus  var.  Lindheimeri,  93. 
ptisillus  var.  oblongifolius,  84. 
pygmaus,  61. 


Davis:    RANUNCULI  OF  NORTH  AMERICA. 


507 


pygmccus  var.  Sabinii,  61. 
radicans  C.  A.  Meyer,  67. 
radicans  Regel,  71. 
ranunculinus,  =  C. 
Rattani,  38. 
recurvatus  Bong,  37. 
recurvatus  Poir,  36. 
recurvatus  Schlecht,  38. 
recurvatus  var.  Nelsoni,  38. 
regulosus,  30. 
reports,  26. 

repens  var.  hispidus,  8. 
repens  var.  macranthus,  22. 
repens  var.  Marilandicus,  16. 
reptans,  89. 

reptans  \av.jiliformis,  89. 
reptans  var  Gormani,  89. 
reptans  var.  intermedius,  90. 
reptans  var.  strigulosus,  90. 
rhomboideus,  48. 
Sab  in  ii,  61 . 
saluginosus,  =  ?  O. 
samolifolius,  86. 
saniculceformis,  36. 
Sardous,  9. 
saxicola,  46. 
sceleratus,  33. 

sceleratus  var.  multifidus,  34. 
Schlechtendalii,  12,  38. 


septentrionalis,  1 2 . 

speciosus,  17. 

stagnatalis,  =  B.  i. 

stolonifer,  87. 

subaffinis^  49. 

subalpinus,  23. 

subsagittatus  var.  subaffinis,  49. 

Suksdorjii,  47. 

sulphztreus,  66. 

tenellus,  37. 

tenellus  var.  Lyalli,  37. 

tomentosus  Poir.,  12. 

tomentosiis  Spreng.,  36. 

trachyspermus  Ell.,  3. 

trachyspenmis  Engelm.,*  92. 

trachyspermus    var.    Lindheim- 

eri,  93. 
trachyspermus      var.       angusti- 

folius,  92. 

trichophyllus,  =  B.  3. 
tridentatus,  =  O. 
trifolius,  7. 
triternatus,  59. 

Turner  i,  39. 

Unalaschensis,  90. 
uncinatus,  56. 
unguiculatus,  78. 
vagans,  88. 
vicinalis,  43. 


XXVII.     A  SYNONYMIC  CONSPECTUS  OF  THE  NA- 
TIVE AND  GARDEN  THALICTRUMS  OF 
NORTH   AMERICA. 


K.  C.  DAVIS. 


The  name  Thalictrwn  (Linn.  Sp.  PI.  545.  1753)  is  prob- 
ably from  Thallo,  to  grow  green,  and  has  reference  to  the 
young  shoots  which  are  of  such  a  bright  green  color. 

This  group  includes  several  forms  which  are  well  suited  for 
the  mixed  border  and  rock  garden,  and  the  robust  forms  are 
well  placed  in  the  wild  garden.  Many  are  very  hardy,  and 
only  the  more  southern  forms  of  those  given  below  are  at  all 
tender.  Thalictrums  are  valued  for  their  healthy  heads  of 
flowers,  contrasted  with  their  handsome  stems  and  leaves  which 
are  often  of  a  purple  cast.  They  may  be  propagated  by  seed 
or  by  division  of  roots  in  early  spring  just  as  growth  begins. 
Any  good  loamy  soil  will  suit  them  if  well  drained. 

The  latest  monograph  of  the  genus  was  in  1885,  by  LeCoyer, 
in  Bull.  Soc.  Roy.  de  Bot.  de  Beige,  where  he  describes  69 
species.  In  1886  Wm.  Trelease  published  a  fine  treatment  of 
"North  American  species  of  Thalictrum"  in  Proc.  Bost.  Soc. 
Nat.  Hist.  23 :  293-304,  in  which  he  recognized  eleven  species 
and  four  varieties  north  of  Mexico.  His  treatment  is  rather 
closely  followed  by  Robinson  in  Gray's  Synop.  Fl.  1895.  Since 
that  time  at  least  ten  new  species  have  been  described — chiefly 
from  Mexico — several  of  which  are  by  J.  N.  Rose  in  Contr.  U. 
S.  Natl.  Herb.  5  :  185,  October  31,  1899.  Only  five  have  been 
introduced  to  our  gardens  from  other  countries. 

Erect  perennial  herbs  :  leaves  ternately  compound  and  de- 
compound ;  stem  leaves  alternate :  flowers  dioecious,  polyg- 
amous, or  perfect  in  some  species,  rather  small,  generally 
greenish-white  or  sometimes  purple  or  yellow,  borne  in  a 
panicle  or  loose  raceme  ;  sepals  4  or  5,  deciduous  ;  petals  want- 
ing ;  stamens  many,  showy  ;  carpels  usually  few,  i-seeded. 

509 


510  MINNESOTA  BOTANICAL  STUDIES. 

KEY  TO  .SPECIES  OF  THALICTRUM. 

A.   Natives  of  United  States  or  introduced. 
B.  Flowers  perfect. 

C.  Filaments  widened  above ;  anthers  ovate,  obtuse. 

D.  Akenes  sessile,  in  a  head,  ovate  oblong petaloideum. 

DD.  Akenes  stalked,  widely  spreading,   straight  along  dorsal 
margin. 

E.   Styles  very  short;  stigma  almost  sessile ,clavatum. 

EE.   Styles  as  long  as  width  of  akene sparcijlorum. 

CC.  Filaments  filiform  ;  anthers  linear,  acute,  or  mucronate. 
D.   Stigma  hastate  or  spurred ;   flower-stem  abruptly  recurved 

near  the  fruit alpinum. 

DD.   Stigma  not  spurred  nor  hastate. 

E.  Fruits  sulcate;   stigma  dilated   on  one  side  of  the  short 

style minus. 

EE.   Fruits  longitudinally  veined ;   stigma  terminal,  minute, 

not  dilated,  style  short glazicum, 

BB.  Flowers  polygamo-direcious. 

C.  Anthers  linear,  mucronate;  filaments  thread-like. 

purpurascens. 
CC.  Anthers  ovate,  obtuse ;  filaments  broadened  above. 

polygamum. 
BBB.  Flowers  direcious,  with  rare  exceptions. 

C.  Filaments  widened  above ;  anthers  ovate,  rather  obtuse. 

aquilegifolium . 

CC.  Filaments  thread-like;  anthers  linear,  acute  or  mucronate. 
D.  Mature  fruits  rather  firm  or  thick-walled,  not  greatly  flat- 
tened, filled  by  the  seed;  leaves  glabrous. 
E.  Blades  of  leaflets  very  thin. 

F.   Roots  tuberous ;  stem  reclining debile. 

FF.   Roots  coarsely  fibrous  ;   stem  erect dioicum. 

.    EE.  Blades  of  leaflets  firm,  veiny  below. 

F.   Styles  clubbed Caulophyttoides. 

FF.   Styles  not  clubbed. 

G.  Akenes  well  stalked coriaceum. 

GG.  Akenes  nearly  sessile -venulosum. 

DD.  Mature  fruits  less  firm,  thin-walled,  2-edged,  either  flat- 
tened or  turgid. 

E.   Leaflets  very  thin occidentale. 

EE.  Leaflets  firm. 

F.   Pistils  5-11  (rarely  13) Fendleri. 

FF.   Pistils  7-20 polycarpum. 

AA.  Natives  of  Mexico,  not  introduced  into  United  States. 
B.  Leaflets  peltate  or  subpeltate. 


Davis:   THALICTRUMS  OF  NORTH  AMERICA.  511 

C.   Akenes  at  least  twice  as  long  as  broad. 

D.  Plant   glaucous    throughout;    akenes    narrowly    elliptical, 

tapering  at  base  or  stalked. 
E.   Divisions  of  leaves  large,  crenations  broad  and  shallow. 

peltatmn. 
EE.  Divisions  of  leaves  small,  with  small  ovate  teeth. 

Jaliscanum. 
DD.   Plant  not   at  all    glaucous;    akenes  subsessile,   one    side 

straight  Cuernavaccmum. 

CC.   Akenes  a  little  longer  than  broad. 

D.   Leaves  once  or  twice  ternate;  leaflets  large,  orbicular. 

Pringlei, 
DD.   Leaves  4  or  5  times  ternate;  leaflets  small,  ovate. 

Guatemalense. 

BB.  Leaflets  subpeltate  on  only  part  of  the  plant .pubigerum. 

BBB.   Leaflets  not  peltate  nor  subpeltate. 
C.   Flowers  perfect. 

D .  Plant  pubescent longistylum. 

DD.   Plant  glabrous Pachucense. 

CC.  Flowers    monoecious     or    polygamous       ( T.     papillosum 

doubtful). 

D.   Plant  glabrous   or  only  glandular,  very  little  or  no  pubes- 
cence. 
E.  Akenes  ribbed,  not  gibbous. 

F.  Leaves  4—5  times  ternate ;  leaflets  i  to  2  inches  across. 

grandiflorum, 
FF.   Leaves  2-3  times  pinnate;  leaflets  small  and  thin. 

Galeottii. 
EE.  Akenes  with  convex  protuberances  on  the  ribs. 

F.  Anthers  mucronate Hernandezii. 

FF.  Anthers  linear,  but  rather  blunt .gibbosum. 

DD.  Plant  pubescent,  hispid  or  hairy. 
E.  Fruits  not  woolly. 

F.   Leaflets  glabrous  above,  glandular-hispid  beneath. 

lanatum. 

FF.   Leaflets  papillose  above;   hairy  veins  prominent  be- 
neath  papillosum. 

EE.  Fruits  woolly tomentettum. 

CCC.  Flowers  dioacious. 

D.   Leaves  pinnately  compound .pinnatum. 

DD.    Leaves  ternately  compound. 

E.  Styles  wanting Madrense. 

EE.   Styles  filiform,  3-4  lines  long .grandifolium. 


512  MINNESOTA    BOTANICAL    STUDIES. 

T.  petaloideum  LINN.  Sp.  PI.  2  ed.  771.     1762. 

Stem  round,  nearly  i  ft.  high,  almost  naked  :  leaves  3-5- 
parted;  leaflets  smooth,  ovate,  entire,  or  3-lobed :  flowers  cor- 
ymbose, perfect ;  sepals  white,  rotund  ;  filaments  pink,  anthers 
yellow :  fruits  ovate-oblong,  striated,  sessile.  June-July. 
Northern  Asia.  Bot.  Cab.  9:  891.  Not  yet  in  trade  lists  but 
well  worthy  of  use  in  gardens  (f).  Lee.  3.^".  75. 

T.  clavatum  DC.  Syst.  i :  171.     1818. 

T.filtyes  TORR.  &  GRAY,  Fl.  i :  38.     1838. 

T.  nudicaule  SCHWEINITZ  ex  T.  &  G.  Fl.  i  :  39.      1838. 

Plant  glabrous,  i  to  2  feet  high,  branching  above  :  leaves  2 
to  3  times  ternate :  leaflets  oval  to  obovate,  rather  large,  thin, 
about  3-lobed  but  variable,  base  variable  :  flowers  perfect,  in  a 
cymose  panicle ;  filaments  spatulate  and  petal-like,  with  short, 
blunt  anthers  :  akenes  widely  spreading  on  weak  stalks  of  nearly 
their  own  length,  obliquely  ovoid,  flattened;  styles  short;  stig- 
ma minute.  May,  June.  Wet  mountain  places,  western  Vir- 
ginia to  Alabama  (f).  Lee.  3.  f.  10. 

T.  sparciflorum  TURCZ.  F.  &  M.  Ind.  Sem.  Petrop.  i :  40. 

1835- 

T.  clavatum  HOOK.  Fl.  i  :  2.      1833,  not  DC. 
T.  Richardsonii  GRAY,  Am.  Journ.   Sci.  42:   17.      1842. 

Stem  erect,  sulcate,  2  to  4  feet  high,  branching,  usually  gla- 
brous :  leaves  triternate,  upper  ones  sessile ;  leaflets  short- 
stalked,  round  or  ovate,  variable  in  size  and  shape  of  base,  round- 
lobed  or  toothed :  flowers  in  leafy  panicles  on  slender  pedicels, 
perfect ;  sepals  obovate,  whitish,  soon  reflexed  ;  filaments  some- 
what widened ;  anthers  very  short :  akenes  short-stalked,  ob- 
liquely obovate,  flattened,  dorsal  margin  straight ;  8-io-nerved  ; 
styles  persistent.  Northern  Asia,  through  Alaska  to  Hudson 
Bay,  in  mountains  in  Colorado  and  southern  California.  Intro- 
duced to  gardens  in  1881.  Lee.  3.  f.  8. 

T.  alpinum  LINN.  Sp.  PI.  545.     1753. 

Stems  smooth,  naked  or  i-leaved,  only  4  to  8  inches  high, 
from  a  scaly  rootstock :  leaves  tufted  at  the  base,  twice  3-5- 
parted  ;  leaflets  coriaceous,  orbicular  or  cuneate  at  the  base, 
lobed,  revolute  :  flowers  in  a  raceme,  perfect ;  sepals  greenish, 
equalling  the  yellow  stamens  :  stigma  linear  ;  akenes  small,  ob- 
liquely obovoid.  Newfoundland  to  Arctic  Alaska,  in  Rockies 


THALICTRUMS    OF    NORTH    AMERICA.  513 

to  southern  Colorado,  Europe,  northern  Asia,  Greenland  and 
Iceland.     Bot.  Mag.  2237.     Lee.  ^.f.  13  (f). 

T.  minus  LINN.  Sp.  PI.  546.     1753. 
T.  saxatile  VILL.  Prosp.  50.     1779. 
T.  purpureum  SCHANG.  in  Pall.  N.  Nord.  Beitr.  6:  42. 

1794. 
T.  saxat His  HORT. 

Stems  round,  sulcate,  i  to  2  feet  high  :  leaflets  variable,  acute 
or  obtusely  lobed,  often  glaucous  :  flowers  drooping  in  loose 
panicles,  perfect ;  sepals  yellow  or  greenish  :  fruit  ovate-oblong, 
sessile,  striated.  Summer.  Europe,  Asia,  northern  Africa. 
A  polymorphous  species  in  the  variation  of  the  leaflets.  Lee. 
5./  2,3,  4. 

Var.  adiantifolium  HORT. 
T.  adriantoides  HORT. 

T.  adiantifolium  BESS,  ex  Eichw.  Skizze  182.      1830. 
Leaflets  resembling  those  of  Adtantum;  a  form  much  used 
and  admired. 

Var.  Kemense  TRELEASE,  1.  c.  300.     1886. 
T.  Kemense  FRIES,  Fl.  Holland,  94.      1817-18. 
T.  minus  var.  elatum  LEG.  1.  c.  283.      1885.     In  part. 
Leaves  thrice-ternate ;  otherwise  much  like  the  type. 

T.  glaucum  DESF.  Thai,  i  ed.  123.     1804. 

T.  speciosum  HORT.  ex  POIR.  Encycl.  5:  315.     1804. 

Stems  erect,  round,  striated,  glaucous,  2  to  5  feet  high  :  leaf- 
lets ovate-orbicular,  3-lobed,  lobes  deeply  toothed  :  flowers  in 
an  erect  panicle,  perfect ;  sepals  and  stamens  yellow :  fruits  4 
to  6,  ovate,  striated,  sessile.  June,  July.  Southern  Europe. 
Lee.  5,/£. 

T.  purpurascens  LINN.  Sp.  PI.  546.     1753. 
T.  rugosum  PURSH.  Fl.  2  :  389.     1814. 
T.  revolutum  DC.  Syst.  i:   173.     1818. 
T.  CornutiMM.  a  HOOK.  Fl.  i:  3.  t.  2.     1833. 
T.  dasycarpum  FISCH.  &  LALL.  Ind.   Sem.  Hort.  Petrop. 

72.     1841. 

T.  purpureum  HORT. 

A  polymorphous  species,  allied  to  T.  polygamum:  stem  3  to 
6  feet  high,  branching  above,  leafy,  pubescent  or  glabrous, 
sometimes  glandular ;  leaflets  larger  than  in  that  type  :  flowers 


514  MINNESOTA    BOTANICAL    STUDIES. 

in  a  long,  loose,  leafy  panicle,  polygamo-dicecious ;  filaments 
narrow,  anthers  rather  long,  taper  pointed :  akenes  slightly 
stalked,  ovoid,  glabrous  or  pubescent  with  6  to  8  longitudinal 
wings;  style  slender,  persistent;  stigma  long  and  narrow, 
Canada  to  Florida,  west  to  the  Rockies.  June,  Aug.  Intro- 
duced 1883. 

Var.  ceriferum  AUSTIN,  Gray  Man.  5  ed.  39.     1867. 
T.  revolutum  LEG.  1.  c.  146.  /.  3,f*  i  - 

This  is  a  variety  with  waxy  glands. 

T.  polygamum  MUHL.  Cat.  54.     1813. 
T.  corynellum  DC.  Syst.  I:   172.      1818. 
T.  Cornuti  var  ,9  HOOK.  1.  c.  3. 
T.  CornutiToKK.  &  GRAY,  Fl.  i:  38.      1838. 
T.  leucostemon  KOCH.  &  BAUCHE,  Ind.  Sem.  Hort.  Berol. 

App.  13.     1854. 

Erect,  3  to  8  or  more  ft.  high,  branching  and  leafy,  smooth  or 
pubescent,  not  glandular  :  leaves  3-4  times  ternate  or  terminally 
pinnate  ;  leaflets  oblong  to  orbicular,  bases  variable,  3-5  apical 
lobes  :  flowers  in  a  long,  leafy  panicle,  polygamo-dioecious ; 
sepals  white  ;  filaments  broadened  when  young  ;  anthers  short: 
akenes  ovoid,  stipitate,  6-8-winged  or  ribbed ;  with  stigmas  as 
long,  which  become  curled.  July-August.  Low  or  wet 
grounds,  Canada  to  Florida,  west  to  Ohio.  Introduced  1881. 
Lee.  2.  f.  12,  T.  pubescens  Pursh,  Fl.  2:  388,  1814,  is 
probably  a  very  pubescent  form  of  this  and  might  be  called 
var.  -pubcscens. 

Var.  macrostylum  ROBINSON,  Syn.  Fl.  i :   17.     1895. 
T,    Cormiti    var.    macrostylum    SHUTTLE,    in    Dist.    PL 

Rugel,  1845-6. 

T.  Cornuti  var.  monostyla  BOT.  ZEIT.  3:   218,  219.     1845. 
T.  macrostylum  SMALL   &   HELLER,   Mem.   Torr.    Club, 

3:   8.     1892. 

Slender;  leaflets  small,  nearly  entire:  fertile  flowers  less 
numerous  and  in  a  more  spreading  panicle  :  akenes  in  a  small, 
dense,  spherical  head.  Mountains  of  western  North  Carolina 
to  Georgia. 

T.  aquilegifolium  LINN.  Sp.  PI.  547.     1753. 

Stem  large,  hollow,  i  to  3  feet  high,  glaucous  :  leaves  once  or 
twice  3-5 -parted  ;  leaflets  stalked  or  the  lateral  ones  nearly  ses- 
sile, slightly  lobed  or  obtusely  toothed,  smooth,  suborbicular : 


Davis:     THALICTRUMS    OF    NORTH    AMERICA.  515 

flowers  in  a  corymbose  panicle,  dioecious  ;  sepals  white ;  sta- 
mens purple  or  white  :  fruit  3-angled,  winged  at  the  angles. 
May-July.  Europe,  northern  Asia.  Bot.  Mag.  1818;  2025 
(as  \tt.formosum).  Garden  47,  p.  357  ;  50,  p.  117.  Lee.  3. 

f-5- 

The  old  name,  T.  Cornuti  L.  Sp.  PI.  545,  may  be  a  syn- 
onym of  this,  and,  if  so,  it  is  the  older  name  being  published 
on  a  preceding  page ;  but  T.  Cornuti  was  described  as  an 
American  plant  which  T.  aquilegifolium  is  not.  As  the  descrip- 
tion and  old  figures  of  T.  Cornuti  L.  do  not  agree  with  any 
American  plant  the  name  may  well  be  dropped,  as  Robinson 
and  DeCandolle  have  suggested.  Those  plants  advertised  as 
T.  Cornuti  are  probably  T.  aquilegifolium, 

T.  debile  BUCKLEY,  Am.  Journ.  Sci.  45:   175.     1843. 

Root  a  cluster  of  fusiform  tubers  :  stem  decumbent,  ^  to  i 
foot  long,  glabrous,  simple  or  branched,  few-leaved  :  leaves 
2  to  3  times  ternate  ;  petioles  long  and  slender ;  leaflets  nearly 
y,  inch  across,  thin,  rotund,  the  3  rounded  lobes  entire  or  again 
lobed,  bases  variable :  flowers  remote,  in  long,  simple  panicles, 
direcious  ;  stamens  often  10,  filaments  little  longer  than  the  an- 
thers :  akenes  2-5,  nearly  sessile,  spreading,  oblong,  not  flat- 
tened, 8-io-ribbed ;  style  minute.  Moist  or  shady  places. 
Georgia  to  Texas.  Lee.  2.f.i  (f). 

Var.  Texanum  GRAY,  Cat.  Coll.  Hall,  PI.  Tex.  3.  1873. 
Name  only. 

Stems  more  rigid  and  erect;  leaflets  smaller,  thicker  and 
nearly  sessile.  A  Texas  form  of  the  above  (t).  Described  in 
Syn.  Fl.  i:  18.  1895. 

T.  dioicum  LINN.  Sp.  PI.  545.     1753. 

T,  Icevigatum  MICHX.  Fl.  I  :  322.     1803. 

T.  Carolinianum  Bosc.  in  DC.  Syst.  I  :   174.      1818. 

Rather  slender,  i  to  2  feet  high,  glabrous  :  leaves  3  to  4  times 
3-parted;  leaflets  thin,  orbicular,  several-lobed  or  revolute, 
bases  variable  :  flowers  in  a  loose,  leafy  panicle  with  slender 
pedicels,  dioecious ;  stamens  much  longer  than  the  greenish 
sepals :  anthers  linear,  obtuse,  exceeding  their  filaments  in 
length  :  akenes  ovoid,  nearly  or  quite  sessile,  longer  than  their 
styles,  with  about  10  longitudinal  grooves.  Early  spring. 
Woods.  Labrador  to  Alabama,  west  to  the  foot  of  the  Rockies. 
Introduced  sometime  before  1891.  Lee.  3.  f.  2,3. 


516  MINNESOTA    BOTANICAL    STUDIES. 

T.    Caulophylloides    SMALL,    Bull.    Torr.    Club,    25:   136. 
1898. 

Plant  glabrous,  2  to  4  feet  high,  from  a  horizontal  rootstock ; 
leaves  on  long  petioles  ;  leaflets  deep  green,  firm,  oval  or  broader 
than  long,  i  to  2  inches  long,  glaucous  beneath  and  with  prom- 
inent nerves,  bases  variable,  apically  3-5-sharp-lobed ;  flowers 
dioecious  :  akenes  elliptic,  3  lines  long,  sharply  ribbed,  con- 
tracted at  the  base  and  stalked,  style  persistent,  clubbed,  ^  the 
length  of  akenes.  Spring  and  summer.  Mountains  of  Ten- 
nessee.— Allied  to  T.  coriaceum,  but  differing  in  the  leaflets  and 
the  shorter  club-shaped  style  (f). 

T.  Coriaceum  SMALL,  Mem.  Torr.  Club,  4:  98.     1893. 
T.  dioicum    var.   coriaceum   BRITTON,  Bull.  Torr.   Club, 
18:  363.      1891. 

Stem  sulcate,  somewhat  branched,  raising  3  to  4  feet  from  the 
yellow  rootstocks  :  leaves  3  to  4  times  ternate,  rather  short-peti- 
oled,  lower  petioles  with  stipule-like  bases  ;  leaflets  coriaceous, 
broadly  obovate,  acutely  toothed  or  lobed  ;  bases  variable  ;  veins 
prominent  on  the  whitish  under  surface :  flowers  in  a  loose 
panicle,  dioecious ;  sepals  and  stamens  whitish  ;  anthers  linear, 
longer  than  the  slender  filaments:  pistillate  flowers  purple; 
akenes  stalked,  oblong-ovoid,  8-io-ribbed  ;  styles  of  less  length, 
persistent.  May-June.  Mountains  of  eastern  Kentucky  into 
Virginia  and  north  Carolina  (f). 

T.  venulosum  TRELEASE,  /.  c.,  302.     1886. 

T.  campestre  GREENE,  Erythea,  4  :   123.      1893. 

(?)  T.    Fendleri].  M.  MACOUN,  Bot.  Gaz.  16 :  285.   1893. 

Allied  to  T.  dioicum :  stem  simple,  erect,  10-20  inches  high, 
glabrous,  glaucous  ;  bearing  2  to  3  long-petioled  leaves  above 
the  base ;  leaves  3  to  4  times  3-parted ;  leaflets  short-stalked, 
rather  firm,  rounded  and  lobed  at  the  apex,  veiny  beneath: 
flowers  in  a  simple  panicle,  dioecious,  small ;  sepals  ovate ; 
stamens  10-20  on  slender  filaments  ;  anthers  oblong,  slender 
pointed  :  akenes  nearly  sessile,  2  lines  long,  ovoid,  tapering  to 
a  straight  beak,  thick-walled  and  2-edged.  South  Dakota, 
westward  and  southward  in  the  mountains.  Introduced  1889. 

T.  occidentale  GRAY,  Proc.  Am.  Acad.  8:  372.     1872. 
T.  dioicum  var.  oxycarpum  TORR.   Bot.    Wilkes  Expe 

212.       1854. 


Davis:    THALICTRUMS  OF  NORTH  AMERICA.  517 

Allied  to  T.  dioicum  which  it  closely  resembles  ;  but  it  is  more 
robust,  taller :  leaves  glandular-puberulent :  akenes  long,  slen- 
der, thin-walled,  2-edged,  ribbed,  not  furrowed.  Introduced 
1881. 

T.  Fendleri  ENGLEM.  ex  Gray  in  Mem.  Am.  Acad.  4:5.    1849. 

A  variable  species  :  plants  i  to  3  feet  high,  rather  stout  and 
leafy  :  leaves  4  to  5  times  pinnatifid,  upper  stem  leaves  sessile; 
leaflets  rather  firm,  ovate  to  orbicular,  usually  with  many  shal- 
low rounded  or  acuminate  lobes,  bases  variable  :  flowers  dioeci- 
ous, in  rather  compact  panicles ;  stamens  many,  anthers  long  : 
akenes  nearly  sessile,  obliquely  ovate,  flattened,  3  to  4  ribs  on 
each  face.  July-Aug.  Western  Texas  to  Montana.  Intro- 
duced 1881.  Lee.  i.f.  9. 

Var.  Wrightii  TRELEASE,  1.  c.  304.    1886. 

T.    Wrightii  GRAY  PI.  Wright  2:  7.     1852. 
The  upper  leaves   petioled ;  leaflets  smaller,  puberulent  be- 
low :    akenes    plump,  sigmoid,   reticulated.     Aug. -Sept.     Dry 
regions.     New    Mexico,    southern    Arizona    into    Chihuahua. 
Lee.  2./.  8. 

Var.  platycarpum  TRELEASE,  1.  c.  304.     1886. 
T.  plalycarpum  GREENE,  Pitt.  1 :   166.     1888. 
T.  hcspermm  GREENE,  Pitt.  2  :   24.      1889. 
Inflorescence  sparsely  glandular-puberulent :  leaflets  like  the 
type:  akenes  flat,  erect,  dilated,  the  veins  mostly  longitudinal- 
Central  and  southern  California. 

T.  polycarpum  WATSON,  Proc.  Am.  Acad.  14 :  288.      1879. 
T.  Fendleri  var.  polycarpum  TORR.  Pac.  R.  Rep.  4:  61. 

1853. 
T.  Fendleri  BREW.  &  WATS.  Bot.  Calif.  1 :  4.     1876.     In 

part. 
T.  ccestum  GREENE,  Fl.  Francis,  309.     1892. 

Allied  to  T.  Fendleri:  glabrous  throughout:  leaflets  long- 
petioled :  flowers  dioecious  in  rather  close  panicles :  akenes 
larger  in  a  dense  globose  head,  stalked,  obovoid,  turgid  taper- 
ing into  reflexed  styles.  Summer.  Sandy  streams,  California 
to  Columbia  river.  Introduced  1881.  Lee.  3.  f.  4. 

T.  peltatum  DC.  Prod,  i :  n.     1824. 

Plant  tall,  glabrous  or  glaucous  :  upper  stem  leaves  twice 
ternate :  leaflets  pale,  3  inches  across,  orbicular,  mostly  peltate, 


518  MINNESOTA    BOTANICAL    STUDIES. 

apically  lobed  or  crenate:  flowers  polygamous  in  an  open  pan- 
icle :  akenes  flattened,  obliquely  oblong,  being  nearly  straight 
on  edge,  base  tapering  but  sessile,  both  faces  2-3-veined ;  styles 
y^  inch  long,  rather  persistent.  August.  Morelos,  south  of 
Mexico  City.  Rose,  1.  c.  186,  redescribes  and  figures  this, 
plate  21.  He  concludes  that  DeCandolle's  type  may  have  been 
found  in  the  same  region.  Type  in  U.  S.  Nat.  Herb.  7448, 
distributed  as  T.  Pringlei  (f). 

T.  Jaliscanum  ROSE,  1.  c.  187.     1899. 

Stems  tall,  glabrous  and  glaucous ;  upper  leaves  ternate,  the 
leaflets  peltate,  orbicular,  6  to  10  toothed,  glabrous  :  inflores- 
cence a  large  open  panicle :  carpels  narrowly  elliptical,  some- 
what cuneate  at  base,  strongly  nerved.  Quoted  from  Rose  by 
whom  it  was  first  collected  on  tableland  in  northeastern  Jalisco. 
Differs  from  T.  -peltatum  in  its  small  leaflets  with  small  rounded 
teeth  (f). 

T.  Cuernavacanum  ROSE,  1.  c.  187.     1899. 

About  2  feet  high,  branching  above,  somewhat  pubescent, 
never  glaucous:  leaves  twice  ternate;  leaflets  roundish,  i  inch 
across,  palmate,  broadly  crenate  :  inflorescence  an  open  panicle ; 
flowers  perfect ;  anthers  linear ;  akenes  2  lines  long,  narrowed 
at  both  ends,  subsessile,  one  side  straight,  3~4-ribbed ;  styles 
long.  Morelos,  south  of  Mexico  City  (f). 

T.  Pringlei  WATS.  Proc.  Am.  Acad.  25:  141.     1890. 
T.  -pubigerum  PRINGLE,  ex  Rose,  1.  c.  187.      1899. 

About  2  feet  high,  glabrous  :  leaves  i  to  2  times  ternate  ;  leaf- 
lets usually  peltate,  suborbicular,  yz  to  2  inches  across,  coarsely 
5~9-toothed,  not  glandular :  inflorescence  an  open  panicle,  with 
slender  nodding  pedicels  ;  flower  polygamo-dioecious  :  anthers 
linear,  long  apiculate  :  akenes  compressed,  semi-ovate,  straight 
on  one  side,  6-8-ribbed,  2  lines  long;  styles  long,  somewhat 
persistent.  June-August.  Near  the  capital  of  Jalisco,  and  the 
coast  slope  of  the  same  state  (f). 

Var  reticulatum  ROSE,  1.  c.  188.     1899. 

A  lower,  somewhat  pubescent  form  :  leaflets  peltate,  entire 
or  3-5 -angled,  dark  green  above,  strongly  net-veined:  flowers 
in  a  narrow  panicle ;  peduncles  nodding  in  fruit.  Western 
foothills  of  Tepic  Territory  (f).  The  type  3372  in  U.  S.  Natl. 
Herb,  is  a  form  of  the  same  variety  with  the  leaflets  shallow- 
round-lobed,  and  some  of  them  only  subpeltate. 


Davis:   THALICTRUMS  OF  NORTH  AMERICA.  519 

T.  Guatemalense  ROSE  &  C.  DC.,  Contr.  Natl.  Herb.  5 :  188. 
1899. 

Stems  about  2  feet  high  ;  slender,  branched,  somewhat  hairy  : 
leaves  4  to  5  times  ternate ;  leaflets  small,  ovate,  peltate,  some- 
what roughened,  strongly  veined  below  :  akenes  turgid,  hardlv 
2  lines  long.  Guatemala  (f). 

T.  pubigerum  BENTH.  PI.  Hartw.  285.     1839-57. 

Plant  rather  tall,  nearly  simple,  glabrous  or  pubescent,  finely 
striated  ;  leaves  2  to  4  times  pinnate  ;  leaflets  distant,  their  stalks 
stipuled,  often  ovate,  sometimes  subpeltate,  cordate  or  roundish 
at  base ;  summit  3-toothed,  often  with  other  smaller  teeth  : 
flowers  monoecious  or  polygamous,  reddish ;  anthers  linear, 
mucronate :  akenes  stipitate,  glabrous,  flattened,  obliquely 
ovate,  reticulately  veined,  protuberiferous,  reflexed ;  style  with- 
ering. Summer.  West  central  Vera  Cruz  (f). 

T.  longistylum  DC.  Syst.  i:  171.     1818. 

Plant  tall,  sparsely  pubescent  even  to  the  fruits :  leaflets 
roundly  lobed  and  toothed  outwardly,  pubescent  beneath  :  flow- 
ers perfect,  anthers  slender,  pointed  :  fruits  flattened  a  little, 
reticulately  ribbed,  straight  along  one  side;  styles  slender, 
longer  than  the  body  before  becoming  broken.  August.  Moist 
banks,  Sierra  de  las  Cruces,  Mexico,  10,000  feet.  Also  South 
America  (f). 

T.  Pachucense  ROSE,  1.  c.  188.     1899. 

Delicate  glabrous  plant,  8  to  12  inches  high;  roots  fibrous: 
leaves  only  3  to  4  inches  long,  mostly  basal,  3  times  ternate  ; 
leaflets  2  to  4  lines  long,  broad  or  narrow,  bases  variable  :  flowers 
perfect,  on  erect  pedicels  which  become  bent  in  fruit ;  sepals 
purplish  ;  anthers  narrow,  apiculate  :  ovaries  oblong  ;  style  long 
and  slender.  Open  woods.  High  altitudes.  Southern  Hi- 
dalgo (f). 

T.  grandiflorum  ROSE,  1.  c.  188.     1899. 

T.  grandifolium  ROSE,  1.  c.  143.  1897.  Not  Wats. 
Stems  5  to  8  feet  high,  glabrous  :  leaves  i  to  2  feet  long,  4  to  5 
times  ternate ;  main  petiole  short  with  long  dilating  stipules  ; 
leaflets  stalked,  large,  nearly  orbicular,  i  to  2  inches  across,  often 
cordate  at  the  base,  3  to  7  roundish  lobes,  sometimes  a  little 
hairy  on  under  veins  :  flowers  in  a  large,  nearly  naked  panicle, 


520  MINNESOTA    BOTANICAL    STUDIES. 

polygamous  ;  filaments  slender  ;  anthers  linear  :  akenes  flattened, 
strongly  nerved,  style  persistent.  Morelos,  south  of  Mexico 
City  (f). 

T.  Galeottii  LEC.  1.  c.  24:  131.  t.  2.f.  6.     1885. 

Rather  tall  and  simple,  glabrous,  2-3-pinnate ;  stipules 
ample ;  leaflets  small,  thin,  ovate  to  obovate,  toothed  or  lobed 
above,  glabrous:  panicles  rather  leafy;  flowers  small,  whitish, 
monoecious  or  polygamous ;  sepals  slightly  dentate ;  anthers 
linear,  somewhat  obtuse:  pistils  5  to  n  :  akenes  nearly  sessile, 
compressed,  glabrous,  semi-ovate,  veined,  widely  spreading ; 
styles  long,  slender,  withering.  September-October.  Moun- 
tains of  central  Vera  Cruz  (f). 

T.  Hernandezii  TAUSCH.  in  Presl.  Reg.  Haenk.  2  :  69.    1835. 

Stem  3-5  feet  high ;  glandulous,  leafy,  branching ;  leaves 
2  to  3  times  5-parted  ;  leaflets  large,  often  subsessile,  variable  in 
outline,  usually  oval,  3-lobed  or  sharp  toothed  above ;  under 
side  glandular:  flowers  in  a  conical  panicle,  monoecious  or 
polygamous ;  sepals  4,  greenish  ;  anthers  linear,  mucronate : 
akenes  5-7,  sessile  or  stipitate,  obliquely  ovate,  compressed? 
spreading,  irregularly  ribbed  and  protuberiferous  ;  styles  long, 
slender,  somewhat  persistent.  June-August.  Southeastern 
Mexico,  Oaxaca,  etc.  Lee.  2.f.  2  (f). 

T.  gibbosum  LEG.  1.  c.  24:   132.  t.  2.f.  7.     1885. 

Tall,  erect,  simple,  or  branched,  glabrous  :  leaves  2  to  3  times 
pinnate,  petiole  stipuled ;  leaflets  small,  thin,  oval,  stalked,  3 
sharp  teeth  above,  often  other  small  ones  :  panicles  slightly 
leafy:  flowers  monoecious  or  polygamous,  small,  greenish; 
sepals  feebly  dentate  ;  anthers  linear,  usually  blunt ;  pistils  4-5  : 
akenes  stalked,  flattened,  tapering  above  and  below,  glabrous, 
strongly  ribbed,  reticulated  and  provided  with  protuberances, 
widely  spreading ;  style  long  and  slender,  withering.  Sept.- 
Oct.  Mountains  of  western  Oaxaca  (f). 

T.  lanatum  LEG.  in  Bull.   Soc.  Roy.  Bot.    Belg.    16 :    226. 

1877. 

Rather  tall  and  leafy,  hispid  :  leaves  2  to  3  times  pinnate,  very 
short  petioled  or  sessile  ;  leaflets  variable  in  size  and  form,  often 
orbicular  or  obovate,  cordate  or  rounded  at  base,  firm,  glandular- 
hispid  beneath,  short-stalked,  tridentate  at  summit,  often  with 


Davis:   THALICTUUMS  OF  NORTH  AMERICA.  521 

other  smaller  teeth  :  panicles  many-flowered,  monoecious  or 
polygamous  ;  sepals  whitish  ;  anthers  linear,  mucronate  :  akenes 
5  to  7,  sessile  or  nearly  so,  spreading,  reticulately  ribbed,  glan- 
dulous;  styles  long  and  filiform,  rather  persistent.  June-Aug. 
Oaxaca,  southeastern  Mexico  (f).  Closely  allied  to  T.  Her- 
vandezii,  but  differing  in  being  glandular  hispid,  and  having  no 
convex  protuberances  on  the  akenes. 

T.  papillosum  ROSE,  1.  c.  189.     1899. 

Low,  hairy :  leaves  small,  3  times  ternate  ;  leaflets  roundish, 
often  cordate  at  base,  somewhat  3-lobed,  papillose  above,  hairy, 
veins  prominent  beneath  :  panicles  short ;  pedicels  becoming  re- 
flexed  in  fruit :  akenes  i  line  long,  few-ribbed.  Northern  Ja- 
lisco and  western  Zacatecas  (f).  Fruit  much  shorter  than  in  T. 
lanatum. 

T.  tomentellum  ROBINSON  &  SEAT.  Proc.  Am.  Acad.  28 : 
103.  1893. 

Stem  striate,  glandular,  finely  and  densely  pubescent  through- 
out :  leaves  3  times  pinnate  on  petioles  i  to  2  inches  long  ;  leaflets 
suborbicular,  subcordate,  shallowly  3-lobed ;  the  lobes  rounded, 
often  with  2  to  3  rounded  teeth  :  flowers  in  a  pyramidal  subnaked 
panicle,  polygamo-dicecious ;  pedicels  becoming  reflexed  in 
fruit :  sepals  2  lines  long  ;  anthers  setiform  at  tip  :  carpels  about 
10,  scarcely  stipulate,  woolly,  roughly  reticulated,  acuminate ; 
style  very  long,  filiform,  often  deciduous.  July.  About  Lake 
Patzcuaro,  Michoacan  (f). 

T.  pinnatum  WATS.  Proc.  Am.  Acad.  23:   267.     1888. 

Hardly  2  feet  high,  glabrous  and  glaucous,  slender :  root 
fascicled,  tubero-fibrous  :  leaves  lanceolate  in  outline,  2^  inches 
long  or  less,  very  shortly  petioled,  pinnate  with  about  7  (or 
fewer)  pairs  of  divisions,  the  lower  divisions  ternate,  with  small 
lobed  leaflets,  the  upper  reduced  to  a  single  3-lobed  leaflet : 
flowers  dioecious  ;  sepals  of  the  fertile  flowers  very  small ;  stig- 
mas short  and  rather  thick  :  akenes  ovate,  about  one  line  long, 
undulately  ribbed,  the  oval  seed  filling  the  cavity.  September. 
Pine  plains,  east  base  Sierra  Madre,  Chihuahua.  Description 
from  the  original  (f). 

T.  Madrense  ROSE,  1.  c.  188.     1899. 

Glabrous,  slender,  i  foot  or  less  high,  from  a  cluster  of  thick- 
ened roots  :  leaves  small,  sessile,  once  or  twice  ternate  :  leaflets 


522  MINNESOTA    BOTANICAL    STUDIES. 

mostly  3-toothed  or  lobed :  flowers  dioecious  (?);  fertile  flowers 
often  axillary  and  single ;  styles  wanting ;  stigma  short  and 
thickened  :  akenes  with  strong,  undulate  ribs.  Quoted  from  Dr. 
Rose,  who  first  collected  it  in  southern  Durango  and  northern 
Tepic  (t). 

T.  grandifolium  WATS.  Proc.  Am.  Acad.  23:  267.      1888. 

Tall,  usually  glabrous  :  leaves  3  to  4  ternate,  petiolate,  with  di- 
lated stipules;  leaflets  i  to  2^  inches  long,  obliquely  rounded, 
often  cordate,  or  upper  ones  cuneate  at  base,  obtusely  lobed, 
veins  prominent  beneath  with  a  few  scattered,  short,  stout,  curved 
hairs  :  panicles  spreading  and  somewhat  leafy-bracteate  ;  flowers 
nodding,  dioecious  :  akenes  semicircular,  beaked  by  the  short, 
stout  base  of  the  long  filiform  style  (3  to  4  lines),  compressed, 
faintly  and  irregularly  nerved :  seed  flattened-subovate,  filling 
the  cavity.  October.  Under  cliffs  of  Sierra  Madre,  Chi- 
huahua (f). 

T.  Wrightii  GRAY,  occurs  in  both  Mexico  and  New  Mexico. 
It  is  placed  in  this  arrangement  as  a  variety  of  T.  Fendleri, 
which  see. 

THALICTRUM   INDEX. 

alpinum  L.  Cuernavacanum  Rose. 

aquilegifolium  L.  dasycarpiim    Fisch.    &.    Lall.  = 

ccesium  Greene  =  polycarptim.  ptirpurascens. 

campestre    Greene  =  venulosum.  debilc  Buckley. 

Carolinianum   Bosc.  =  dioicum.  deb  He  var.   Texanum  Gray. 

cazilophylloides  Small.  Delavayi  Franc. 

clavatum  DC.  dioicum  L. 

clavatum  Hook.  =  sparciflorum.  dioicum   var.    coriacium    Britton 

coriacium  Small.  ==  coriacium. 

Cornuti  L.,  see  aquilegifolium.  Fendleri  Engelm. 

Cornuttvax.  monostyla  Bot.  Zeit.  Fendleri  var.  platycarpum  Tre- 

=  polygamum  var.  lease. 

Cornuti    var.      macrostylum  Fendleri  Macoun.  =  venulosian. 

Shuttlew.  =  polygamum  var.  Fendleri  Brew.  &  Wats.  =  poly- 

Cornuti    var.     a    Hook.  =  ptir-  carpum. 

purascens.  Fendleri  var.   Wrz'g-fitiiTrelcase. 

Cornuti    var.     /3    Hook.  =  poly-  Fendleri  var.  polycarpum  Torr 

gamum.  =  polycarpum. 

Cornuti  T.  &  G.  =  polygamum.  filipes  T.  &  G.  =  clavatum. 

corynellum  DC.  —polygamum.  Galeottii  Lee. 


Da-vis:    THAI.ICTRUMS  OF  NORTH  AMERICA. 


523 


gibbosuin  Lee. 

glaiicum  Desf. 

grandiflorium  Rose. 

grandi folium  Rose  =  above. 

grandifolium  Wats. 
GantcmalcHsc  Rose  &  C.DC. 

Hcruandczii  Tausch. 

hespcrium  Greene=ttnd/crivar. 

Jaliscanum  Rose. 

Kcmense  Fries.  =  minus  var. 

favigatum  Michx.  =  dioicum. 

taxation  Lee. 

mttcastemon  Koch.  &  Bauche.  = 
polygamum. 

lo  ngistylu  m  DC. 

macrostyhtm   Rob.    var.    of  poly- 
gam it  m, 

maci-ostylum    Small    &  Heller  = 
polygamum  var. 

Madrcnsc  Rose. 

minus  L. 

minus  var.  Kemense  Trelease. 

minus  var.  elattim  Lee.  =  minus 


midicaule  Schw.  =  clavatum. 
occidentale  Gray. 
Pachuccnse  Rose. 
papillosum  Rose. 
pcltatum  DC. 


pctaloidcum  L. 

pinnatum  Wats. 

platycarptim   Greene  =  Fendleri 
var. 

Pringlei  Wats. 

Pringlei  var.  reticulatum  Rose. 

polycarpicm  Wats. 

polygamum  Muhl. 

pzibcscens     Pursh  =  polygamum 
var. 

pubigerum  Benth. 

purpurascens  L. 

purpureum     Hort.  =  purpuras- 
cens. 

purpureum  Schang.  =  minus. 

revolutum    DC.  =  prirpurascens. 

revolutum    Lee.  =  purpurascens 
var. 

Richardsonii       Gray  =   sparci- 
Jlorum. 

rugosum    Pursh  ^  purpurascens. 

saxatile  Vill.  =  minus. 

saxatilis  Hort.  =  mimis. 

sparcijlorum  Turcz. 

speciosum  Poir.  =  glaucum. 

tomentellum  Rob.  &  Seat. 

venulosum  Trelease. 
Wrightii  Gray  =  Fendleri  var. 


NOTE. — The  mark  (|)  indicates  that  the  species  or  variety 
has  not  yet  been  introduced  to  the  American  trade.  Citations 
after  descriptions  are  mostly  to  pictures.  '.'  Lee."  refers  to  Le- 
Coyer's  monograph. 


XXVIII.    SOME  PRELIMINARY  OBSERVATIONS  ON 
DICTYOPHORA  RAVENELII  BURT. 


C.    S.    SCOFIELD. 


The  name  Dictyo-phora  was  first  applied  by  Desvaux  in  1809 
to  a  plant  bearing  a  netted  veil  or  indusium,  and  the  genus  so 
named  was  later  included  under  the  general  family  Phalloidese, 
established  by  Fries  in  1823.  The  family  was  given  thorough 
systematic  arrangement  by  Dr.  Ed.  Fischer  *  in  1888,  and  in 
1896  Dr.  E.  A.  Burt  f  published  a  systematic  account  of  the  ten 
known  North  American  species  under  six  genera. 

The  development  of  the  sporophore  has  been  especially 
studied  in  plants  of  this  family,  and  this  process  has  been  de- 
scribed for  many  of  the  species.  In  the  present  paper  less  at- 
tention has  been  given  to  this  particular  feature  of  the  life 
history,  not  that  it  is  less  interesting',  but  because  in  some  of  the 
stages  it  is  not  dissimilar  to  other  species  that  have  already  been 
well  described  and  figured. 

Collection  of  material. — The  material  for  the  study  of  Dic- 
tyo-phora raveneln  was  collected  about  September  25,  1899. 
It  was  found  on  low  moist  ground  in  rather  dense  woods  near 
the  west  shore  of  Lake  Calhoun,  Minneapolis,  Minnesota. 
The  mycelium  of  the  plant  was  more  or  less  abundant  over  an 
area  of  five  or  six  square  yards,  and  the  sporophores  seemed  to 
occur  over  most  or  all  of  this  extent.  The  period  of  fruiting  is 
evidently  long,  for  photographs  of  the  mature  plants  were  made 
at  least  a  month  before  the  material  was  collected,  and  at  the 
time  of  collection  sporophores  in  nearly  all  stages  of  develop- 
ment were  abundant. 

Two  collections  of  material  were  made :  that  of  the  first  col- 
lection was  put  directly  into  thirty  per  cent,  alcohol  and  after- 
ward passed  gradually  into  ninety  per  cent.  ;  while  that  of  the 

*  Saccardo,  Syl.  Fung.  7  :  1888. 
tBot.  Gaz.  22:  1896. 

525 


526  MINNESOTA    BOTANICAL    STUDIES. 

second  collection  was  placed  in  a  one  per  cent,  solution  of 
chromic  acid,  from  which  after  twenty-four  hours  it  was  trans- 
ferred to  water  and  after  thorough  washing  was  carried  by  easy 
stages  into  seventy  per  cent,  alcohol. 

Methods. — The  material  for  study  was,  with  a  few  excep- 
tions, dehydrated,  imbedded  in  paraffin,  and  cut  with  a  Minot 
microtome,  carried  down  to  fifty  per  cent,  alcohol,  stained  in  a 
fifty  per  cent,  alcohol  saturated  solution  of  Bismarck  brown, 
transferred  into  pure  xylol  and  permanently  mounted  in  Canada 
balsam.  Some  of  the  small  portions  of  the  mycelium  and 
younger  stages  in  the  development  of  the  sporophore  were  first 
stained  in  toto,  and  either  mounted  directly  in  formalin  water 
and  sealed  or  transferred  to  paraffin  and  cut  and  stained  again 
if  necessary.  The  pre-staining  method  proved  very  effectual 
and  was  of  great  help  in  guarding  against  the  loss  of  very  small 
bodies,  and  aided  in  the  imbedding  process.  Numerous  other 
staining  methods  were  tried,  but  none  gave  as  good  result  for 
structural  study  as  the  one  outlined. 

The  vegetative  tract  consists  of  a  complex  weft  of  mycelial 
strands,  which  vary  in  size  from  something  less  than  one-tenth 
of  a  millimeter  in  diameter  up  to  two  millimeters  or  more. 
The  complexity  of  the  weft  is  greatly  augmented  by  the  copi- 
ous branching  of  the  strands  and  not  uncommonly  crossing 
strands  become  more  or  less  fused  together.  Some  of  the 
larger  strands  have  a  length  of  one  meter  or  more,  and  often 
continue  with  unvarying  diameter  for  forty  or  fifty  centimeters. 
The  larger  proportion  of  the  mycelium  is  found  near  the 
surface  of  the  soil  where  it  is  covered  with  leaf  mould  and  may 
be  found  to  some  extent  in  the  leaf  mould  itself.  Some  of  it, 
however,  runs  to  a  considerable  depth  in  the  soil,  but  without 
diminishing  in  size  or  ending  there  as  would  a  root  of  a  higher 
plant.  Invariably,  strands  found  at  the  greatest  depth  of  twenty 
to  thirty  centimeters  could  be  traced  to  the  surface  in  both  di- 
rections. Branching  seems  to  be  less  frequent  on  the  strands 
found  deep  in  the  soil,  and  it  was  not  possible  to  locate  in  any 
case  what  seemed  to  be  the  definite  center  of  growth. 

Each  mycelial  strand  is  composed  of  two  general  areas :  the 
central  and  the  peripheral.  In  the  very  small  threads  the  cen- 
tral area  (Fig.  9,  .Z?)  consists  of  a  few  large  hypha?,  very  long 
in  proportion  to  their  diameters,  aud  without  very  definitely 
marked  cross-septa.  Their  general  direction  is,  of  course, 


Scofidd  :     DICTYOPHORA    RAVEXELII    BURT.  527 

along  the  strand,  but  they  are  more  or  less  twisted  about  each 
other,  very  much  as  are  the  threads  which  make  up  a  strand  of 
yarn. 

The  peripheral  area  (Fig.  9,  c]  is  composed  of  loosely  inter- 
twining hyphas,  much  smaller  and  more  profusely  branched 
than  the  central  hyphas,  and  extending  out  somewhat  into  the 
surrounding  soil  (Fig.  8).  They  seem  to  resemble  very  much 
the  root  hairs  on  the  roots  of  higher  plants.  It  seems  quite 
probable  that  the  hyphas  of  the  peripheral  areas  of  the  smaller 
strands  function  as  the  absorptive  area  of  the  plant,  while  the 
larger  central  hyphas  act  as  conduction  paths. 

In  the  larger  strands  the  peripheral  hyphas  occupy  a  rela- 
tively smaller  part  of  the  strand  and  seem  to  abandon  their  ab- 
sorptive, to  assume  more  of  a  cortical  function,  being  reduced 
in  the  very  large  strands  to  a  smooth  disorganized  coating. 
The  central  hyphas  by  their  habit  of  twisting  about  each  other 
make  it  difficult  to  determine  their  method  of  growth  and 
branching,  for  they  do  not  continue  long  enough  in  the  plane 
of  the  section  to  be  studied  with  ease,  and  in  no  case  were  defi- 
nite cross-septa  noted  although  they  doubtless  exist. 

Upon  the  mycelium  are  borne  two  distinct  kinds  of  bodies  : 
(i)  the  reproductive  body,  and  (2)  what  it  has  seemed  best  to 
call  a  storage  body  or  "  tuber."  The  latter  will  be  considered 
first. 

The  tuber  makes  its  appearance  as  a  slight  enlargement  of  a 
mycelial  strand,  and  in  the  early  stages  of  its  development  seems 
to  be  merely  the  result  of  rapid  growth  of  the  peripheral 
hyphas.  There  seems  to  be  little  regularity  in  the  size  or  shape 
of  the  tubers  and  even  less  in  regard  to  their  place  of  occur- 
rence upon  the  mycelium.  In  Fig.  iconeof  the  larger  tubers  is 
shown,  natural  size,  and  upon  a  connecting  strand  is  shown  at 
"«  "  the  base  of  an  old  sporophore.  The  strand  bearing  this 
tuber  seems  to  have  been  more  or  less  branched  and  the  tuber 
is  lobed  to  some  extent  to  follow  the  branching.  The  tuber  is 
made  up  of  very  closely  woven  hyphas  which  are  much  dis- 
torted, evidently  by  being  packed  full  of  somewhat  granular 
material.  In  general  structure  it  appears  homogeneous  except 
the  region  of  the  strand  upon  which  it  is  borne,  where  the 
hyphas  seemed  to  leave  the  strand  to  some  extent  and  mingle 
with  those  of  the  tuber,  but  not  so  much  so  that  the  direction  of 
the  strand  cannot  be  clearly  followed  throughout.  A  section 


528  MINNESOTA    BOTANICAL    STUDIES. 

of  a  young  tuber  is  diagrammatically  shown  in  Fig.  u.  The 
cell  contents  of  the  hyphae  of  the  tuber  respond  very  neatly  to 
a  test  for  glycogen  given  by  Dr.  L.  Errera,*  and  it  seems  very 
probable  that  this  substance  is  a  very  large,  if  not  the  chief 
constituent  of  the  cells.  Errera's  test  is  made  with  a  reagent 
composed  of  450  parts  of  water,  three  parts  of  KI  and  one 
part  of  iodine,  and  he  designates  it  as  "  solution  lodee  au  ^-g-." 
According  to  him,  material  possessing  glycogen  when  stained 
in  this  solution  or  when  mounted  in  a  drop  of  it  takes  on  a  red- 
dish brown  color,  which  disappears  in  a  temperature  of  50-60° 
C.,  but  returns  upon  recooling.  Some  of  the  tests  made  on 
Dictyoflwra  tubers  were  with  material  taken  from  70  per  cent. 
alcohol  sectioned,  transferred  to  water  and  mounted  directly  in 
a  drop  of  the  reagent. 

In  other  instances  sections  that  had  been  machine  cut,  stained 
in  Bismarck  brown,  and  mounted  in  Canada  balsam  were  soaked 
in  xylol  to  remove  the  cover  glass  and  balsam,  carried  through 
alcohol  to  water,  stained  for  a  moment  in  Errera's  mixture,  and 
then  mounted  in  water.  In  every  case  the  reaction  was  sharp 
in  all  particulars. 

Assuming  that  Errera's  test  is  a  correct  one,  and  there  ap- 
pears no  good  reason  for  doubting  it,  it  is  evident  that  a  large 
portion  at  least  of  the  cell  contents  of  the  tuber  is  glycogen. 
Zopf ,  Burt  f  and  others  have  associated  the  presence  of  gly- 
cogen in  fungi  with  the  immediate  need  of  the  plant  for  rapid 
growth,  but  there  is  at  least  a  possibility  that  this  reserve  food 
supply  in  the  tuber  is  in  some  way  connected  with  the  economy 
of  the  plant  in  reproducing  itself  vegetatively  the  following 
season.  Or  it  may  be  that  there  exists  a  direct  connection  be- 
tween the  tuber  and  the  rapidly  developing  sporophore,  though 
there  is  no  evidence  that  any  of  the  supply  of  glycogen  has  been 
exhausted  from  any  of  the  tubers  collected  or  found  upon  the 
vegetative  tract.  If  it  is  found  upon  further  investigation  that 
these  tubers  are  connected  with  vegetative  reproduction  and 
that  by  means  of  them  it  is  possible  to  artificially  propagate  the 
plant,  it  will  be  of  great  advantage  in  the  study  of  the  younger 
stages  of  development  of  the  reproductive  area.  Hitherto  the 
rare  occurrence  of  the  plant  and  its  allies  has  made  the  study 
extremely  difficult. 

*  Leo  Errera,  Sur  le  Gljcogene  ches  les  Basidiomycetes,  1885. 
t  Bot.  Gaz.  24  :     1897. 


Scoficld  :     DICTYOPHORA    RAVENELII    HURT.  529 

The  reproductive  area  usually  occurs  on  a  branch  strand  of 
the  mycelium.  The  length  of  this  branch  varies  with  the  dis- 
tance of  the  main  strand  below  the  surface  of  the  soil.  In  some 
instances  this  branch  is  so  short  that  the  sporophore  seems  ses- 
sile upon  the  main  strand.  Often  the  sporophore-bearing  branch 
and  the  strand  from  which  it  comes  are  very  small  (Fig.  8), 
and  the  main  strand  here  seems  to  diminish  but  little  in  size 
after  giving  off  the  branch.  Both  the  branch  and  the  strand, 
however,  increase  in  size  as  the  sporophore  develops.  In  all 
cases  the  development  of  the  sporophore  takes  place  very  close 
to  the  surface  of  the  soil  so  that  upon  nearing  maturity  it  pushes 
partially  above  the  surface  before  the  rupturing  of  the  volva 
and  the  elongation  of  the  stipe  takes  place.  A  number  of 
these  nearly  mature  sporophores  are  shown  in  an  accompanying 
plate.  Before  taking  up  the  development  of  the  sporophore  a 
brief  description  of  the  mature  organ  will  be  given  to  explain 
the  parts  and  to  define  the  terms  used. 

The  mature  sporophore  (Fig.  7)  is  8— 10  cm.  high  and  con- 
sists of  a  base  B,  volva  F1,  F2,  F3,  stipe  S,  indusium  /,  pi- 
leus  P  and  gleba  G.  The  base  may  be  considered  for  the  pres- 
ent as  a  part  of  the  volva,  although  structurally  and  develop- 
mentally  it  doubtless  belongs  to  the  same  area  as  the  stipe. 
The  volva  is  slightly  pinkish  and  consists  of  more  or  less  defi- 
nitely organized  outer  and  inner  layers  with  a  disorganized 
milky-gelatinous  layer  between.  The  stipe  is  hollow,  2-3  cm. 
in  diameter,  dirty  white,  tapering  at  each  end,  with  walls  com- 
posed of  several  layers  of  chambers  and  passing  with  the  pileus 
into  a  thick,  white,  recurved  collar  at  the  distal  end.  The 
border  of  the  collar  is  entire,  not  convoluted  as  in  some  species. 

The  indusium  or  veil  is  membraneous  and  not  of  definite 
structure.  It  is  attached  at  the  point  of  union  of  the  stipe  and 
pileus  and  also  to  the  volva  near  the  base  (Fig.  7,  /).  It  rup- 
tures irregularly  upon  the  elongation  of  the  stipe  and  may  break 
from  near  either  the  point  of  attachment,  or  partially  from  both, 
and  hang  about  the  stipe  as  is  shown  in  the  first  cut.  Most 
commonly,  however,  it  seems  to  break  from  near  its  attachment 
to  the  volva  and  hang  between  the  pileus  and  stipe. 

The  pileus  is  conic-campanulate,  2.5-3.5  cm.  long  and  2-3 
mm.  thick,  dirty  white  and  with  an  entire,  slightly  recurved 
margin.  It  is  composed  of  closely  folded  layers  of  pseudo-paren- 
chymatous  tissue,  which  give  to  its  surface  a  finely  wrinkled  or 
granulate  appearance. 


530  MINNESOTA    BOTANICAL    STUDIES. 

The  gleba  is  thin,  slightly  shorter  than  the  pileus,  very  dark 
olive  green  and  much  firmer  in  texture  and  more  persistent  than 
is  common  with  the  other  members  of  the  genus ;  deliquescing 
slowly  in  dry  weather  and  without  so  much  of  the  foetid  odor 
common  to  the  plants  of  this  class. 

The  description  of  the  development  of  sporophore  must  be- 
gin with  the  youngest  stage  found,  although  manifestly  a  com- 
plete description  should  start  rather  with  the  activity,  nuclear  or 
cytologic,  that  takes  place  before  the  spore-bearing  branch  is 
formed. 

In  the  youngest  stage  found  (Fig.  i),  the  sporophore  was 
about  .4  mm.  in  diameter  and  borne  upon  a  branch  about  .1 
mm.  in  diameter.  The  young  sporophore  consisted  of  but  two 
distinguishable  areas  ;  the  central  (Fig.  i,  r]  and  the  peripheral 
(Fig.  i,  V),  the  chief  difference  being  that  the  hyphae  of  the 
central  area  were  somewhat  larger  and  took  a  much  deeper 
stain  than  those  of  the  other.  The  two  areas  of  the  strand 
seemed  to  be  continued  into  the  sporophore  with  a  slight  in- 
crease in  the  proportional  space  occupied  by  the  outer  one.  The 
line  between  them  is  not  as  clearly  marked  as  in  the  strand,  the 
hyphag  being  closely  anastomosed.  A  detail  of  the  structure 
of  this  stage  is  shown  in  Fig.  12.  Much  time  and  ingenuity 
was  spent  in  attempting  to  determine  the  condition  in  the  strand 
just  previous  to  the  formation  of  the  sporophore.  It  seems 
evident  that  one  must  look  for  the  starting  point  in  the  main 
strand  or  at  least  in  the  very  young  branch. 

There  seems  to  be  good  reason  for  believing  that  some  cell 
fusion  may  take  place  in  the  strand  previous  to  the  giving  off 
of  the  sporophore  branch.  In  Fig.  8  is  shown  a  small  mycelial 
strand  with  a  branch  "a"  leading  to  a  very  small  sporophore. 
Near  the  middle  of  this  strand  is  shown  one  hypha  much  more 
prominent  than  the  rest,  so  much  so  that  it  may  readily  be  seen 
through  the  surrounding  tissue,  and  by  careful  focusing  its 
course  may  be  traced  for  some  distance  either  side  of  the  place 
of  branching.  It  is  difficult  to  see  through  the  tissue,  and  still 
more  difficult  to  get  sections  to  show  whether  or  not  an  actual 
fusion  has  taken  place,  which  has  given  rise  to  a  new  body. 
Evidently  a  fusion  of  some  kind  may  have  occurred,  and,  in 
the  reaction  following,  one  of  the  hyphas  may  have  come  to  be 
of  a  slightly  different  nature,  for  the  single  hypha  is  not  par- 
ticularly prominent  except  near  the  branching  point.  The  uni- 


Scofidd :     DICTYOPIIORA    RAVENELII    HURT.  531 

form  presence  of  this  prominent  hypha  would  suggest  the  idea 
that  it  is  intimately  concerned  with  the  formation  of  the  sporo- 
phore  while  also  perhaps  evidence  that  a  fusion  has  taken  place 
at  this  point  may  be  seen  in  the  knotted  condition  of  the  hypha. 
This  gives  rise  to  the  thought  that  a  cell  fusion  at  this  point 
may  have  initiated  all  the  resulting  activities.  In  Fig.  9  is 
shown  a  section  of  a  small  strand  at  the  branching  point  and 
the  supposed  evidence  of  fusion  is  here  very  clear.  That  the 
peculiar  deep-staining  ability  of  this  prominent  hypha  is  conse- 
quent to  such  a  fusion  is  indicated  by  the  fact  that  the  hypha 
cannot  be  traced  along  the  strand  any  great  distance  from  this 
point. 

It  is  unfortunate  that  the  technique  of  the  material  is  not 
sufficiently  developed  to  make  possible  a  study  of  the  nuclear 
phenomena  at  this  point,  for  clearly  the  complete  solution  of 
this  problem  must  lie  in  the  study  of  the  nuclear  processes. 

Whatever  action  takes  place  here  is  a  matter  of  considerable 
importance  in  the  life  history  of  the  plant,  for  the  subsequent 
differentiation  of  the  hyphal  tissue  is  very  complex. 

The  first  marked  step  in  the  differentiation  of  the  sporophore 
is  shown  in  Fig.  2.  The  gelatinization  of  the  area  between 
the  outer  and  inner  layers  of  the  volva  is  shown  at  V2.  There 
is  a  somewhat  indefinite  integument  formed  about  the  whole 
body  by  the  breaking  down  of  the  hyphae  at  the  surface.  The 
hyphse  lying  in  the  area  of  the  stipe  S  also  begin  to  be  promi- 
nent and  tissue  of  much  the  same  nature  extends  out  like 
an  umbrella  from  the  top  of  the  stipe  area.  This  is  evidently 
brought  about  by  the  apical  growth  of  the  large  hyphae  shown  in 
Fig.  i.  Being  limited  by  the  denser  hyphae  of  the  periphery, 
they  take  a  downward  direction.  This  tissue  "/>"  gives  rise 
later  to  the  pileus  and  gleba  and  there  remains  between  this  and 
the  stipe  a  tissue  of  the  same  structure  as  that  of  the  volva. 
The  tissue  of  the  base  "^"  is  similar  in  structure  to  that  of  the 
stipe,  but  closer  in  texture.  The  next  important  stage  is  shown 
in  Fig.  3.  Here  the  different  areas  are  fairly  well  marked. 
The  area  between  pileus  and  stipe  is  distinct,  but  is  composed 
of  very  loosely  woven  hypha?.  It  is  in  direct  connection  with 
the  tissue  of  the  volva,  but  is  nearly  separated  from  it  by  the 
base  ".Z?,"  which  has  extended  considerably. 

There  becomes  evident  at  this  point  an  area  of  less  tension 
near  the  middle  of  this  base  and  just  below  the  stipe.  This  is 


532  MINNESOTA    BOTANICAL    STUDIES. 

connected  by  a  small  pore,  through  the  base,  with  the  partially 
disorganized  tissue  in  the  center  of  the  stipe.  The  stipe  already 
shows  very  slightly  its  chambered  structure  and  the  areas  of  the 
pileus  and  gleba  are  distinguishable. 

In  Fig.  4  the  relative  size  of  the  various  parts  is  shown  to  be 
considerably  modified.  Gelatinization  of  the  middle  area  of  the 
volva  is  nearly  complete.  The  indusium  is  almost  entirely  cut 
off  from  the  volva  and  occupies  a  much  smaller  space  than  in  the 
previous  stage.  The  gleba  is  greatly  enlarged,  and  the  hyme- 
nial  layer  is  beginning  to  show  and  the  pressure  of  the  whole  re- 
ceptaculum  is  beginning  to  be  exerted  upon  the  inner  layer  of 
the  volva.  The  next  stage  as  shown  in  Fig.  5  shows  general 
enlargement  of  the  parts  and  rapid  development.  Just  how  this 
enlargement  takes  place  is  not  easy  to  understand.  Certainly 
it  is  not  altogether  due  to  enlargement  of  existing  hyphce  for 
excepting  in  the  stipe  and  pileus  actual  measurement  of  the 
cells  in  the  various  stages  show  slight  differences  in  the  sizes  of 
individual  cells,  so  that  enlargement  must  be  very  largely  due  to 
apical  growth  and  branching. 

The  development  as  shown  from  Figs.  2  to  5  must  go  on  very 
rapidly,  for  comparatively  few  sporophores  in  these  stages  were 
found.  In  Fig.  6  is  shown  the  sporophore  practically  mature. 
The  tissue  in  the  middle  of  the  stipe  JTis  almost  completely  dis- 
organized, showing  the  wall  of  the  other  side  of  the  stipe  in  one 
or  two  places.  The  walls  of  the  stipe  are  fully  developed,  but 
the  cells  of  the  walls  of  the  chambers  are  closely  compressed, 
especially  at  the  angles.  The  indusium  is  reduced  to  a  thin 
layer  adhering  closely  to  the  sides.  The  inner  layer  of  the 
volva  is  drawn  very  tightly  over  the  gleba  and  is  pressed  against 
the  outer  layer  at  the  tip.  The  gleba  is  completely  developed 
and  the  spores  are  nearly  or  quite  formed. 

Fig.  7  shows  the  mature  plant  as  previously  described.  The 
parts  of  it  may  now  be  described  in  detail.  The  base  B  is 
made  up  of  small  but  entire  hyphas  closely  interwoven  and 
shown  in  detailed  structure  in  Fig.  19.  The  base  forms  a  cup 
which  loosely  contains  the  base  of  the  stipe  to  which  it  is  at- 
tached only  slightly,  just  about  the  pore  which  connects  the 
hollow  of  the  base  with  the  hollow  of  the  stipe.  The  tissue 
of  the  central  area  of  the  strand  is  in  direct  connection  with 
the  tissue  of  the  base  and  the  peripheral  area  of  the  strand  at 
this  time  greatly  reduced  leads  directly  into  the  coating  of  the 
base  and  the  outer  layer  of  the  volva. 


Scofidd :     DICTYOPHORA    RAVENELII    BURT.  533 

The  volva  having  been  so  tightly  compressed  before  rupturing, 
has  its  two  layers  so  close  together  that  they  might  readily  be 
mistaken  for  one,  and  the  detailed  structure  is  hard  to  recog- 
nize. The  outer  layer  is  hardened  and  the  interstices  between 
the  hyphae  are  filled  with  gummy  material.  The  inner  layer 
has  its  hyphoi  lying  for  the  most  part  in  the  direction  of  the  re- 
cent strain  and  connects  by  a  thin  layer  with  the  indusium. 

The  stipe  is  made  up  of  several  rows  of  irregular  chambers 
as  shown  in  Fig.  13  in  cross  section.  Some  of  these  chambers 
open  to  the  outside,  but  none  of  them  to  the  middle  of  the  stipe. 
The  chamber  walls,  one  of  which  is  shown  in  detail  in  Fig.  17, 
are  made  up  of  much  distended  hyphae  which  look  in  section 
like  the  parenchymatous  cells  of  higher  plants.  The  chambers 
contain  filmy  remnants  of  disorganized  tissue.  The  remnant  of 
the  tissue  in  the  hollow  of  the  stipe  X,  hangs  usually  from  the 
apex  of  the  stipe  or  some  of  it  may  remain  attached  at  the  base. 
The  chambers  of  the  stipe  walls  become  smaller  toward  either 
end  and  at  the  apex  the  wall  passes  into  the  recurved  collar 
where  the  contents  of  the  chambers,  though  somewhat  disor- 
ganized, are  not  absorbed. 

The  indusium  which  has  been  the  cause  of  the  trouble  in 
classification  is  hardly  to  be  considered  the  true  indusium  com- 
mon to  the  genuine  members  of  the  genus  Dictyophora.  It  is 
not  a  definitely  organized  structure,  but  rather  the  remnant  of  a 
portion  of  the  tissues  of  the  periphery  of  the  young  sporophore 
caught  between  the  pileus  and  the  base.  Penzig*  in  describing 
Ithypkallus  tennis,  speaks  as  follows  :  "  Eine  andere  bemerken- 
swerthe  Erscheinung  bei  Ithy-pallus  tennis  ist  das  Auftreten 
einer  Art  von  Indusium  zwischen  dem  Hute  und  dem  oberen 
Theile  des  Stieles.  Auch  Ed.  Fischer  (1.  c.,  p.  22)  kurz  die 
Andesenheit  einer  Haut,  welche  er  als  '  Rest  der  stiel  und  Hute 
trennenden  Primordialgewedes '  auffast.  Dieses  Gedielde  ist 
nicht  in  alien  Individuen  gleich  ausgedildet :  einmal  nur  als 
ausserest,  feines,  durchsichtiges  Hautchen,  andere  Male  aber  als 
zienlich  derbe,  compacte  membran,  welche  den  Stiel  kragen- 
artig  oder  fast  glockenformig  umgiebt.  Ihere  structur  ist  nicht 
pseudoparenchymatisch  ;  vielmehr  ist  sie  aus  eng  verflochtenen, 
cylindrischen  Hyphen  zusammengesetz." 

Burtf  in  describing  this  plant  writes  :   "  This  species  has  been 

*Ann.  Jard.  Bot.  de  Buitenzorg,  2d  Ser.     Vol.  I.,  part  2. 
tBot.  Gaz.  22.     1896. 


534  MINNESOTA    BOTANICAL    STUDIES. 

placed  in  the  genus  Dictyophora  on  account  of  its  having  a 
persistent  membrane  hanging  about  the  angle  between  the  pileus 
and  the  stipe.  This  membrane  is  composed  of  the  same  tissue, 
the  intermediate  tissue  A  of  my  figures  ;  which  gives  rise  to  the 
veil  in  D.  duplicata.  Differentiation  of  this  tissue  does  not 
advance  in  D.  ravenelii  to  the  final  stage  of  making  this  mem- 
brane pseudoparenchyma,  or  is  this  final  stage  reached  in  the 
case  of  hyphas  composing  the  pileus  in  /.  impudtcus  and  in  D. 
duplicata,  yet  no  one  would  hesitate  on  that  ground  to  use  the 
term  pileus  in  connection  with  those  species.  It  seems  best  to 
apply  the  term  veil  to  this  membrane  in  D.  ravenelii  which 
looks  like  a  veil,  has  the  position  of  a  veil,  is  composed  of  a 
tissue  forming  the  veil  in  other  species  and  is  likely  to  be  re- 
garded as  a  veil  without  question  by  every  botanist  meeting  this 
fungus  for  the  first  time  and  attempting  its  determination." 

Burt's  interpretation  of  this  structure  seems  hardly  the  best 
one.  The  membrane  as  shown  in  detail  in  Fig.  15  bears  no 
resemblance  whatever  to  the  tissue  of  the  stipe  and  pileus  shown 
in  detail  with  the  same  enlargement  in  Figs.  14  and  16.  The 
attachment  of  this  membrane  at  the  base  of  the  volva  and  the 
fact  that  it  ruptures  irregularly,  are  both  strong  reasons  for  not 
considering  it  as  a  true  veil  or  indusium.  It  would  seem  better 
to  regard  it  rather  as  tissue  which  in  other  species  of  both 
Itkyphallus  and  Dictyophora,  is  completely  disorganized — with 
the  exception  possibly  of  /.  tennis,  mentioned  by  Penzig,  where 
also  a  similar  structure  occurs.  The  presence  of  this  tissue,  al- 
though noted  by  the  earlier  writers,  was  not  considered  of  im- 
portance enough  to  exclude  the  plant  from  the  genus  in  which 
its  other  characteristics  certainly  placed  it. 

The  pileus  is  composed  of  tissue  very  similar  in  structure  to 
that  of  the  stipe,  except  that  the  walls  are  closely  folded  and 
the  tissue  of  the  chambers  is  not  so  completely  disorganized. 
The  structure  of  one  of  the  folds  is  shown  in  Fig.  14  while  a 
longitudinal  section  showing  the  relative  position  of  the  walls  is 
shown  in  Fig.  18.  From  a  surface  view  the  pileus  has  a  finely 
wrinkled  or  granulated  appearance.  It  is  firmly  attached  to  the 
apex  of  the  stipe  just  below  the  collar.  The  line  of  demarka- 
tion  between  the  pileus  and  the  stipe  at  the  point  of  attachment 
is  not  easy  to  make  out.  In  fact  at  the  point  of  union  the  tissue 
of  the  three  areas,  stipe,  pileus  and  collar  is  homogeneous.  It 
is  close  within  the  axis  of  the  stipe  and  pileus  that  the  so-called 


Scofidd  :     DICTYOPHORA    RAVENELII    BURT.  536 

inclusium  is  attached  and  often  it  clings  so  closely  to  the  inner 
surface  of  the  pileus  as  to  be  mistaken  for  a  portion  of  that 
structure. 

The  gleba  is  much  more  persistent  than  in  most  forms  of  the 
family,  maintaining  itself  for  some  hours  after  the  elongation  of 
the  stipe.  Its  structure  at  this  stage  is  very  indefinite.  Slight 
traces  of  the  hymenium  may  be  found,  but  for  the  most  part  it 
consists  of  a  disorganized  tissue  containing  masses  of  spores 
scattered  about,  held  by  the  surrounding  substance.  The  spores 
are  very  small,  1.5-2.5  mikrons  in  diameter,  somewhat  oblong 
and  greenish  black.  In  order  to  show  the  structure  of  the 
gleba  the  drawings  for  Figs.  16  and  17  were  made  from  a 
young  stage  of  the  sporophore  such  as  is  shown  in  Fig.  6. 

In  conclusion  the  results  of  the  study  so  far  made  upon  this 
plant  seem  to  suggest  the  following  points  : 

1.  The  mycelium  of  the  plant  is  of  considerable  structural 
importance  and  deserves  further  attention. 

2.  There  are  borne  upon  the  mycelium  certain  organs  which 
seem  to  function  as  storage  places  for  reserve  material. 

3.  There  is  in  the  young  mycelial  threads  very  good  evidence 
of  the  occurrence  of  cell  fusion  previous  to,  or  in  intimate  con- 
nection with  the  formation  of  the  sporophore. 

4.  The  indusium  of  this  plant  cannot  be  considered  homol- 
ogous with  the  indusium  of  true  members  of  the  genus  Dictyo- 
phora;  but  is  rather  the  persisting  remnant  of  tissue  which  is 
completely  broken  down  in  most  other  plants  of  this  order. 

EXPLANATION  OF  PLATES. 

PLATE  XXIX. — Field  view  of  undeveloped  sporophores,  from 
photograph  by  C.  J.  Hibbard. 

PLATE  XXX. — Field  view  showing  mature  sporophore,  from 
photograph  by  C.  J.  Hibbard. 

PLATE  XXXI. — Structure  and  development  of  /.  ravenelii. 

i.  A  very  young  sporophore ;    V,  volva;  7?,  receptaculum.      x  5°- 

2-7.  Development  of  the  sporophore ;  B,  base ;  V,  the  outer  layer 
of  the  volva;  K2,  the  middle  layer  of  the  volva;  K3,  the  inner  layer 
of  the  volva  ;  /,  indusium  ;  S,  stipe  ;  P,  pileus  ;  G,  gleba  ;  C,  collar, 
and  A'",  tissue  remnant  within  the  stipe. 

8.  Small  mycelial  strand  with  branch  "a"  leading  to  young  sporo- 
phore. x  50. 


536  MINNESOTA    BOTANICAL    STUDIES. 

9.  Section  of  small   strand   at  point   of    branching,  showing    large 
hy pha  at  "  a  "  ;  central  area  B ;  peripheral  area  C.      x  i  oo. 

10.  Tuber  with  connecting  mycelium  and  base  of  an  old  sporo- 
phore  at  A.     Natural  size. 

1 1 .  Section  through  a  small  tuber    showing    continuous    mycelial 
strand.      X  5- 

12.  Detail    of    portion   of    young   sporophore    shown    in   Fig.    i. 
X  500. 

13.  Diagrammatic  view  of  section  of  stipe.      x  5. 

14.  Detail  of  a  fold  of  pileus  tissue.      x  500. 

15.  Detail  of  portion  of  indusium  showing  large   drops  of  gelatine 
at  hyphal  ends.      X  500. 

1 6.  Portions  of  hy  menial  layer  in  immature  sporophore.      x  500 

17.  Portion  of  wall  of  chamber  of  stipe,      x  500. 

18.  Diagrammatic  view  of  pileus  and  gleba  immature.      x  15. 

19.  Detail  of  portion  of  tissue  of  the  base  of  the  sporophore.      x 
500. 


VOL.  II. 


MI.NNKSOTA  BOTANICAL 


PART  IV. 


VOL.  II. 


MIXXKSOTA    lioTAxu  \i.  STUDIES. 


PART   I  V 


VOL.  II 


MlNNESC 


:AJL  STUDIES. 


PART  IV. 


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XXIX.    A    PRELIMINARY    LIST    OF   MINNESOTA 
UREDINE^E. 


E.  M.  FREEMAN. 


The  following  list  comprises  the  Uredinese  collected  in  Min- 
nesota up  to  the  present  time,  by  the  Botanical  Survey  Staff. 
The  materials  are  taken  from  the  collections  cited  in  my  pre- 
liminary list  of  Minnesota  Erysipheae.*  In  addition  to  these 
Dr.  L.  H.  Pammel  has  made  numerous  collections  at  Hokah 
and  other  points  in  southeastern  Minnesota.  These  have  not 
been  included  in  the  present  report  but  may  be  found  in  Tre- 
lease's  Parasitic  Fungi  of  Wisconsin. f  Puccinia  anemones- 
virginiantB  is  the  only  species  collected  by  Dr.  Pammel  in  Min- 
nesota which  has  not  been  collected  elsewhere  in  the  State. 

No  representatives  of  the  Endophyllacese  or  of  the  Schizo- 
sporaceae  have  yet  been  found  in  Minnesota.  Of  the  Melamp- 
soraceag  five  genera  with  seven  species  are  reported,  viz: 
Chrysomyxa  i  species,  Cronartium  I,  Cpleosporium  i,  Me- 
lampsora  3,  Calyptospora  i  ;  of  the  Pucciniaceae  seven  genera 
with  62  species:  Uromyces  14,  Puccinia  39,  Gymnoconia  i, 
Uropyxis  i,  Gymnosporangium  4,  Phragmidium  4,  Triphrag- 
mtum  i  ;  of  isolated  ^Ecidia  (including  Peridermid)  30 ;  of 
isolated  Uredo  2. 

On  May  n,  1900,  there  was  collected  in  Mille  Lacs 
county  a  very  large  witches'  broom  on  a  white  pine.  The 
broom  measures  fully  9  feet  across.  The  distortion  of  the 
branches  is  very  pronounced  and  the  leaves  of  the  broom  are 
considerably  smaller  than  the  normal.  The  cause  of  the  for- 
mation cannot  at  present  be  positively  ascertained.  There  are 
no  indications  that  the  branches  contain  an  abundant  mycelium 
and  the  material  was  collected  early  in  the  spring  before  ascidia 
had  time  to  form.  So  far  as  I  am  aware  no  authentic  record  of 
a  witches'  broom  upon  pines  caused  by  a  fungus  parasite  exists. 


*Minn.  Bot.  Stud.  2< :  417.     1900. 

f  Trans.  Wise.  Acad.  Sci.  A.  and  L.  6:  1884. 

537 


538  MINNESOTA    BOTANICAL    STUDIES. 

In  Sargent's  Sylva*  is  a  statement  that  pines  are  sometimes  sub- 
ject to  the  distortions  known  as  witches'  brooms.  Dr.  Farlow 
writes,  however,  that  this  statement  was  based  on  a  reported 
witches'  broom  on  Pinus  -ponderosa  from  Montana  and  that 
further  study  of  the  material  demonstrated  that  the  distortions 
were  not  typical  witches'  brooms,  nor  were  they  caused  by  a 
fungus  parasite.  No  definite  statement  as  to  the  cause  of  the 
broom  of  white  pine  collected  in  Minnesota  can  be  made  until 
older  material  is  obtained  and  a  more  detailed  description  is 
therefore  deferred. 

DESCRIPTION  OF  PLATE  XXXII. 

Witches'  broom  on  Pinus  strobus,  Mille  Lacs  county,  Minn.  After 
photograph  by  R.  S.  Mackintosh,  May,  1900. 

I.  MELAMPSORACE^E. 

Chrysomyxa  UNGER. 

One  species  of  this  genus  has  been  found.  Common  on 
Pyrolas  throughout  the  State. 

i.  C.  pirolatum  (KOERN.)  WINT.  Die  Pilze  i1 :   250.     1884. 
On  leaves  of : 

Pyrola  rotundifolia  L.  :   Goodhue,  (II)  Je.  1893,  Ballard  ; 

Freeborn,  (II)  My.  1891,  Sheldon  5964  and  5963. 
Pyrola  elliptica.  NUTT.  :  Aitken,    (II)  Je.    1892,    Sheldon 
2101  ;  Houston,  (II)  Je.  1899,  Lyon  98;  Mille  Lacs,  (II) 
My.  1900,  H.  B.  Carey  and  Freeman  560. 
Pyrola  secunda  L.  :   St.  Louis,   (II)  Jy.   1886,  Hoi  way  27. 
[  Urcdo  pyrolcs  (Gm.)  Wint.j 

Cronartium  FRIES. 

Not  yet  collected  by  the  survey  staff,  but  one  species  has  been 
reported  by  Seymour. 

i.  C.  asclepiadeum  (WILLD.)  FR.  Obs.    Myc.  i:   220.     1815. 
Var.  quercuum  B.  &  C. 

Crow  Wing,  (III)  Ag.  1884,  Seymour.     (Economic  Fungi. 
A.  B.  Seymour  and  F.  S.  Earle,  No.  215.) 

Coleosporium  LEVEILLE. 

One  species  found.  Very  abundant  throughout  the  State. 
Uredospore  form  most  abundant  and  conspicuous.  Only  one 
collection  of  the  teleutospores  has  been  made. 

*Svlva  of  N.  A.  ii  :  12. 


Freeman:    AIINXKSOTA  UKEDINE./E.  539 

i.  C.~§onchi-arvensis  (P.)  WINT.  Die  Pilze  i1  :  247.     1884. 
On  leaves  of  : 

Solidago  serotina  AIT.:  Lincoln,  (II)  Ag.  1891,  Sheldon 
1506;  Houston,  (II)  Ag.  1899,  Lyon  360. 

Solidago  canademis  L.  :  Traverse,  (II)  S.  1893,  Sheldon 
7080. 

Solidago  flexicaulis  L.  :  Case,  (II)  Ag.  1893,  Anderson 
702. 

Solidago  sp.  indet.  :  Chicago,  (II)  S.  1891,  Sheldon  4261  ; 
Lincoln,  (II)  Ag.  1891,  Sheldon  1420;  Hennepin,  (II) 
O.  1898,  Freeman;  Traverse,  (II)  S.  1893,  Sheldon 
7308  ;  Houston,  (II)  Ag.  1899,  Lyon  430. 

Laciniaria   sp.    indet.  :    Traverse,   (II)  S.  1893,  Sheldon 


Aster  divaricatus  L.  :  St.  Louis,  (II)  Jy.  1886,  Holway  102. 
Aster  macrophyllus    L.  :    Cass,    (III)   Ag.    1893,    Ballard 

1747. 
Aster  sp.  indet.  :  St.  Louis,  (II)  Jy.  1886,  Holway  165  and 

251;  Winona,  (II)  Jy.  1888,  Holzinger  139;   Houston, 

(II)  Ag.   1899,  Lyon  401   and  398;  Hennepin,  (II)  S. 

1889,  MacMillan;    Hennepin,  (II)  Ag.   1892,  Sheldon 

4122. 
Docllingeria    umb  elicit  a   (DiLL.)    NEES  :     St.    Louis,  Jy. 

1886,  Holway  83. 

Melampsora  CASTAGNE. 

Three  species.  The  uredo  forms  of  those  species  growing 
on  Populus  and  Salix  are  especially  abundant.  All  three 
species  are  Hemi-melampsoras. 

1.  M.  epilobii  (P.)  FCKL.  Sym.  Myc.  44.     1869. 
On  leaves  of  : 

Efolobium  color  atum  MUHL.  :  -  ,  (II)  S.  1893,  Sheldon 
6147  ;  St.  Louis,  (II)  Jy.  1886,  Holway  35  and  89. 

Epilobium  linear  e  MUHL.  :  St.  Louis,  (II)  Jy.  1886,  Hol- 
way 49. 

Epilobium  sp.  indet.  :  Waseca,  (II)  Je.  1891,  Sheldon  357. 

2.  M.  populina  (jAcc^)  LEV.  Ann.  Sci.  Nat.  III.  8  :  375.    1847. 
On  leaves  of  : 

Popilus  tremuloides  MICHX.  :  Otter  Tail,  (II)  Ag.  1892, 
Sheldon  3890  ;  St.  Louis,  (II)  Jy.  1886,  Holway  198. 


540  MINNESOTA    BOTANICAL    STUDIES. 

Po-puhis  deltoides  MARSH.  :  Lincoln,  (II)  Ag.  1891,.  Shel- 
don 1573  ;  Hennepin,  (II)  S.  1899,  Lyon ;  Winona,  (II). 
S.  1888,  Holzinger;  Hennepin,  (II)  S.  1889,  Mac- 
Millan. 

3.  M.  salicis-capreae  (P.)  WINT.  Die  Pilze  i2:  239.     1884. 
On  leaves  of : 

Salix  discolor  MUHL.  :   St.    Louis,    (II)  Jy.  1886,   Holway 

101. 
Salix  myrtilloides  L.  :  St.   Louis,   (II)  Jy.   1886,   Holway 

166. 

Salix  sp.  indet.  :  Hennepin,  (II)  S.  1890,  MacMillan ; 
Brown,  (II)  Jy.  1891,  Sheldon  1087  and  995  ;  Blue 
Earth,  (II)  Jy.  1891,  Sheldon  478;  Mille  Lacs,  (II)  Jy. 
1892,  Sheldon  2978;  Hennepin,  (II)  O.  1892,  Sheldon 
4126. 

Calyptospora  J.  KUHN. 

The  well-known  species  on  the  mountain  cranberry  has  been 
collected  only  in  one  locality. 

i.  C.  goeppertiana  KUHN.  Hedw.  8:  81.     1869. 

On  Vaccinium  vitis-idcea  L.  :  Cooke,  (III)  Jy.  1899,  Mac- 
Millan ;  Cooke,  (III)  Jy.  1900,  Mrs.  C.  J.  Hibbard. 

II.  PUCCINIACE^E. 
Uromyces  LINK. 

Fourteen  species  of  Uromyces  have  been  found  in  the  State. 
Seven  of  these  are  found  upon  plants  of  the  Pulse  family.  Very 
common  also  are  those  species  found  upon  Euphorbia^  Arts&ma 
and  Polygonum.  Four  species  belong  to  the  Hemiuromyccs^ 
seven  to  the  Euuromyces  and  of  the  remaining  three,  the  life 
histories  are  incomplete.  The  Euuromycetes  are  all  autcecious. 

A.    AUT-EUUROMYCES. 

i.  U.  fabae  (P.)  DE  BARY,  Ann.  Sci.  Nat.  IV.  20:  76.  1863. 
Uromyces  -polymorj)hus  Pk.  *  differs  from  this  species  only  in 
greater  variability  of  the  teleutospore  form.  The  pedicel  is  no 
criterion.  I  have  therefore  included  forms  on  Lathyrus  and 
Vicia  under  U.fabce  although  exhibiting  considerable  variation 
in  spore  form. 


*  Ellis.     N.  A.  Fungi  no.  1442. 


Freeman :    MINNESOTA  UREDINE,<E.  541 

On  Vicia  linear  is  (NuTT.)  GREENE:  Traverse,  (II)  S. 
1893,  Sheldon  7374. 

Vicia    americana   MUHL.  :  Mille   Lacs,   (II  and    III)   Jy. 

1892,  Sheldon  2926. 

Vicia  sp.  indet.  :  Traverse,  S.  1893,  Sheldon,  (II)  7320 
and  (III)  7256; ,  (III) 1892,  Sheldon  3816  (?). 

Lathyrus  venosus  MUHL.  :  Mille  Lacs,  (II)  Jy.  1892,  Shel- 
don 2940. 

Lathyrus  sp.   indet.  :  ,  (III) I893>  Sheldon  6127 

and  6129. 

2.  U.    appendiculatus    (P.)   LINK.   Berl.    Ges.  Nat.    Freunde 

Mag.  7:  28.     1816. 

On  Strophostyles  helvola  (L.)  BRITTON  :  Houston,  (I)  Je. 
1899,  Lyon  24;  Houston,  (II,  III)  Ag.  1899,  Lyon  389. 

3.  U.  albus  DIET.  &  HOLW.  Hedw.  36:  297.     1897. 
According  to  Dietel  this  is  ^Scidium  album  Clint.  (s£cidt'um 

porosumPk.).  No  experiments  indicating  this  are  cited.  The 
specimen  reported  below  differs  from  Dietel's  description  in 
that  the  teleutospore  sori  are  found  on  the  leaves  (not  on 
the  stem),  and  the  color  of  the  sori  is  not  black  but  dark 
brown. 

This    material    (Holway  25)   was    reported  by    Arthur  *    as 
Uromyces  orobi  (P.)  Wint.  (II),  and  (Holway  14)  as  sEcidium 
porosum. 

On  Vicia  americana  MUHL.  :  St.  Louis,  (II,  III)  Jy.  1886, 
Holway  25  ;  St.  Louis,  Jy.  1886,  Holway  14 ;  Mille 
Lacs,  (I)  Jy.  1892,  Sheldon  2720;  Aitkin,  (I)  Jy.  1892, 
Sheldon  2658. 

4.  U.  trifolii  (ALB.  &  SCHW.)  Wint.  Die  Pilze   I1 :  159.      1884. 

On  Trifolium  re-pens  L.  :  St.  Louis,  (I,  II)  Jy.  1886, 
Holway  34 ;  Hennepin,  (II,  III)  Jy.  1890,  MacMillan ; 
,  (II) 1893,  Sheldon  6098. 

5.  U.   euphorbias  COOKE  &  PECK,  Rep.  N.  Y.  St.  Mus.  Nat. 

Hist.  25:  90.     1873. 

Arthur's  recent  experiment  f  although  too  incomplete  to  be 
conclusive  indicates  strongly  that  Uromyces  euphorbia  is  an  au- 
toecious  Euuromyces. 


*1.  c. 

f  Cultures  of  Uredineae  in  1899.    J-  C-  Arthur. 


542  MINNESOTA    BOTANICAL    STUDIES. 

On  Euphorbia  maculata  L.  :     Mille  Lacs,   (I)  Jy.    1892, 

Sheldon  3136. 
Euphorbia   scrpyllifolia    Pers.  :     Pine,  (II,  III)  Je.    1899, 

Freeman  528;  Hennepin,  (II,  HI)  Jy.  1890,  MacMillan. 
Euphorbia  heterophylla  L.  :      Houston,  (II,  III)  Ag.  1899, 

Lyon  317. 
Euphorbia    marginata    PURSH  :       Renville,    (II,    III)   Jy. 

1891,  Sheldon  957. 

Euphorbia  glyptosperma  ENGELM.  :  Chisago,  (II,  III) 
Ag.  1892,  Taylor  1568^;  Brown,  (II,  III)  Jy.  1891, 
Sheldon  969  and  1170. 

Euphorbia  sp.  indet.  :  Renville,  (II,  III)  1890,  Mac- 
Millan. 

6.  U.  polygon!  (P.)  FCKL.  Symb.  Myc.  64.     1869. 

Of  this  very  common  species  uredospores  and  teleutospores 
only  can  at  present  be  reported.  Some  ascidium  material  on 
what  appeared  to  be  a  Polygonum  was  found  in  1900  in  Wright 
county  (Freeman  698),  but  the  amount  of  material  was  insuffi- 
cient for  accurate  and  positive  determination. 

On  Polygonum  aviculare  L.  :  Winona,  (III)  on  leaves  and 
stem,  N.  and  O.  1893,  Edna  Porter;  St.  Louis,  (II)  Jy. 
1886,  Holway  5,  113  and  116. 

Polygonum  ramosissimum  MICHX.  :  Brown,  (II,  III)  on 
leaves  and  stem,  Jy.  1891,  Sheldon  1048;  Lincoln,  (III) 
Ag.  1891,  Sheldon  1545. 

Polygonum  sp.  indet.  :  Traverse,  (III)  on  leaves  and  stem, 
S.  1893,  Sheldon  7253;  Winona,  (II,  III)  S.  1888, 
Holzinger. 

7.  U.  caladii  (Scnw.)  FARLOW,  Ellis  N.  A.  Fungi,  No.  232. 

1879.     Abundant  wherever  Arisama  is  found. 
On  the  leaves  and  spathe  of : 

Arisama  triphyllum  (L.)ToRR.  :  Blue  Earth,  (I)  Je.  1891, 
Sheldon  114;  Chisago,  (III)  S.  1893,  Sheldon  6309; 
Hennepin,  (I)  My.  1899,  Freeman  308  ;  Ramsey,  (I) 
My.  1899,  Freeman  317  ;  Houston,  (I)  Je.  1899,  Lyon 
97;  Hennepin,  (I)  My.  1899,  MacMillan;  Hennepin, 
(I)  F.  1899,  E.  A.  Cuzner  (in  university  plant  house) ; 
Wright,  (I)  My.  1900,  Freeman  581  ;  Pope,  (III)  Jy. 

1892,  Taylor  916;  Winona,  My.  1889,  Holzinger. 


Freeman:    MINNESOTA  UREDINE/E.  543 

B.   HEMIUROMYCES. 

8.  U.  lespedezse  (Scnw.)  PK.  Ellis,  N.  A.   Fungi,  No.   245. 

1879. 

On  Lcspedeza    capitata    MICHX.  :  Winona,  (III)  S.  1888, 
Holzinger;  Chisago,  (III)  S.  1893,  Ballard  1819. 

9.  U.   hedysari-paniculata  (Scnw.)  FARLOW,    Ellis,   N.   A. 

Fungi,  No.  246.     1879. 
On  Mcibomia  sp.  indet.  :  ,  (III)  1893,  Sheldon  7078. 

10.  U.   terebinth!   (DC.)  WINT.  Die  Pilze  i1 :   147.     1884. 
On  Rhus  radicans  L.  :  Kandiyohi,  (III)  Jy.  1892,  Frost  302. 

11.  U.    caryophyllinus    (SCHRANK.)    SCHROET.    Brand    and 

Rost-Pilz.  Schles.  10.     1872. 

On  Dianthus  caryophyllus  L.  :   Ramsey,  (III)  on  leaves 
and  stem,  Ap.  1900,  Freeman. 

C.  LIFE  HISTORIES  INCOMPLETELY  KNOWN. 

12.  U.   argophyllae  SEYM.    Proc.  Bost.   Soc.   Nat.    Hist.    185. 

1889. 

On  Psoralca  argophylla  PURSH  :    Traverse,  (III)  S.  1893, 
Sheldon  7353;  Lincoln,  (III)   Ag.  1891,  Sheldon    1546. 

T3-  U.  pyriformis  COOKE,  Rep.  N.  Y.  St.  Mus.  Nat.  Hist.  29: 

69.     1878. 

On  Acorus  calamus  L.  :  Carver,   (III)  Je.    1891,  Ballard 
13  ;  Wright,  (III)  My.  1900,  Freeman  636. 

14.  U.  rudbeckiae  ARTH.  &  HOLW.  Bull.  111.  St.  Lab.  Nat. 

Hist.  2:   163.      1885. 
On  Rudbcckia  laciniata  L.  :  ,  (III)  S.  1893,  Sheldon. 

Puccinia  PERSOON. 

Thirty-nine  species  have  been  collected  :  10  Aut-eupuccinia , 
6  Heler-eupuccinia ,  2  Brachypuccinia,  2  Pucciniopsis  ^  7  Hemi- 
puccinia,  i  Micropuccinia,  3  Leptopuccinia,  and  8  with  im- 
perfectly known  life  histories.  Very  abundant  are  those  species 
found  on  Mints,  Helianthus  and  allied  genera,  on  grasses, 
sedges  and  Polygonum.  Those  forms  formerly  included  under 
P.  hieracii  (Schum.)  Mart,  have  been  separated  as  far  as  pos- 
sible according  to  the  recent  researches  of  Jacky.*  The  high 
degree  of  specialization  in  the  habit  of  these  forms  which  has 

*Die  Compositen-bewohnenden  Puccineen  vom  Tjpus  Puccinia  hieracii  und 
deren  Specialisierung.  Bern.  1899. 


544  MINNESOTA    BOTANICAL    STUDIES. 

been  demonstrated  in  these  experiments  emphasizes  the  need 
of  special  research  in  cultures  upon  American  species.  It  is 
possible  to  utilize  Jacky's  results  only  upon  species  common 
to  both  Europe  and  America.  Puccinia  amorphcz  Curt,  upon 
species  of  Amorpha  have  been  retained  under  Schroeter's  genus 
Uropyxis . 

A.   AUT-EUPUCCINIA. 

1.  P.  adoxae  HEDW.  Fl.  Fr.  2:  220.      1815. 

On  Adoxa  moschatellina  L.  :  Winona,  (I)  My.  1889,  Hol- 
zinger. 

2.  P.  calthae  LK.  in  Linne,  Sp.  PI.  62:  79.     1825. 

On  Caltha  palustris  L.  :  St.  Louis,  (II)  Jy.  1886,  Holway 
96. 

3.  P.  convolvuli  (P.)  CAST.  Obs.  i :  16.     1842. 

On  Convolvulus  sepium  L.  :  Winona,  (II,  III)  Ag.  1888, 
Holzinger;  Brown,  (I)  Jy.  1893,  Sheldon  899;  Blue 
Earth,  (I)  Je.  1891 ;  Sheldon  374. 

Convolvulus  spithamceus  L.  :  Winona,  (III)  Ag.  1888,  Hol- 
zinger 6. 

4.  P.  galii  (P.)  SCHWEIN.   Syn.  Fung.  Car.  Sup.  73.      1822. 
On  Galium  asprellum  MICHX.  :   St.  Louis,  Jy.  1886,  Hol- 
way 94. 

Galium  continuum  TORR.  &  GRAY  :  Winona,  Ag.  1888, 
Holzinger  198. 

5.  P.  gentianae  (STRAUSS)  LK.  in  Linne  Sp.  PI.  62:  73.    1825. 
On  Genttana andrewstt  GmsEB.  :  Brown,  (II,  III)Jy.  1891, 

Sheldon. 

Gentiana  puberula  MICHX.  :  Glenwood,  (II,  III)  Ag.  1891, 
Taylor  1179. 

6.  P.  pimpinella  (STRAUSS)  LINK  in  Linne  Sp.  PI.  62:  77. 

1825. 

On  Washingtonia  claytoni  (Micnx.)  BRITTON  :  Houston, 
(II,  III)  Je.  1899,  Lyon  41. 

7.  P.  violae  (SCHUM.)  DC.  Fl.  Fr.  6:  92.     1815. 

On  Viola  canadensis  L.  :  Lake,  (I)  Je.  1893,  Sheldon 
4758- 

Viola  blanda  WILLD.  :  St.  Louis,  (II,  III)  Jy.  1886,  Hol- 
way 131  ;  Lake,  (I)  Je.  1893,  Sheldon  4735  ;  Wright, 
(I)  My.  1900,  Freeman  662. 


Freeman:    MINNESOTA  UREDINE^E.  545 

Viola  blanda  pal  it  str  if  or  mis  A.  GRAY  :  Hennepin,  (I)  My. 
1891,  Sheldon  5961. 

Viola  hlauda  uiiuvna  (LE  CONTE)  B.S.P.  :  Crow  Wing, 
(I)  Je.  1892,  Sheldon  2150;  Mille  Lacs,  (I)  Jy.  1892, 
Sheldon  2705^. 

Viola  scahrinscula  (T.  &  G.)  SCHWEIN.  :  Wright,  (I)  My. 
1900,  Freeman  592. 

Viola  sp.  indet.  :  Hennepin,  (II,  III)  S.  1889,  MacMillan  ; 
Brown,  (III)  Jy.  1891,  Sheldon  845  and  850;  Henne- 
pin, (III)  O.  1892,  Sheldon  4090;  -  -  ,  (III)  S.  1893, 
Sheldon  7242. 

8.  P.  menthae  americana  BURRILL,  Bull.  111.   St.  Lab.  Nat. 
Hist.  2  :   189-191.      1885. 

The  echination,  which  distinguishes  this  form  from  the  Euro- 
pean form,  is  in  almost  all  specimens  most  marked  at  the  apex 
of  the  teleutospores.  Many  teleutospores  are  almost  smooth 
at  the  base.  European  teleutospore  specimens  are  sometimes 
slightly  echinate  at  the  apex. 

On  Monarda  fistulosa  L.  :  Chisago,  Ag.  1883,  Arthur; 
Winona,  (III)  Ag.  1888,  Holzinger  42  ;  Winona,  (II, 
III)  Ag.  1888,  Holzinger;  Hennepin,  (III)  1890,  Mac- 
Millan; Hennepin,  (III)  O.  1893,  Sheldon  4096;  Trav- 
erse, (III)  S.  1893,  Sheldon  7175  ;  Winona,  (III)  S. 
1893,  Edna  Porter;  Hennepin,  (III)  S.  1898,  MacMil- 
lan; Houston,  (III)  Ag.  1899,  Lyon  321. 
Koellia  virginiana  (L.)  MAcM.  :  Winona,  (II,  III)  Ag. 

1888,  Holzinger  164. 

Mentha  canadcnsis  L:  Hennepin,  (II,  III)  Ag.  1883,  Ar- 
thur; St.  Louis,  (II,  III)  Jy.  1886,  Holway  236;  Lin- 
coln, (II,  III)  Ag.  1891,  Sheldon  1419  ;  Ramsey,  (II)  Je. 
1899,  Freeman  458;  Ramsey,  (II,  III)  S.  1898,  Free- 
man. 

Mentha  sp.  indet.  : ,  (II,  III)  1893,  Sheldon  6070  and 

7019;  Traverse,  (II,  III)  S.  1893,  Sheldon  7382. 

9.  P.  tanaceti  DC.  Fl.  Fr.  2:  222.     1815. 

The  Puccinia  on  Helianthus  differs  from  that  on  Tanacetum 
only  in  having  a  slightly  broader  teleutospore.  This  is  espe- 
cially true  of  the  distal  cell.  Culture  experiments  are  necessary 
to  separate  these  forms. 

On  Artemisia  dracunculoides  PURSH  :  Houston,  (II,  III)  Ag. 
1899,  Lyon  391  ;  Traverse,  (III)  S.  1893,  Sheldon  7311. 


546  MINNESOTA    BOTANICAL    STUDIES. 

Heliopis  hclianthoides  (L.)  B.S.P.  :  Winona,  (II,  III)  Jy. 

1888,  Holzinger  204. 
Helianthus  grosse-serratus  MARTENS  :   Lincoln,  (III)  Ag. 

1891,  Sheldon  1544. 
Helianthus  tuberosus  L.  :   Goodhue,    (II,  III)    Ag.    1893, 

Anderson  726. 
Helianthus  giganteus  L.  :   St.  Louis,  (II)  Jy.  1886,  Holway 

133- 

Helianthus  annuus  L.  :  Winona,   (III)  Ag.   1889,  Holzin- 

ger ;    Hennepin,   (II,    III)   O.    1889,    MacMillan  ;    Mc- 

Leod,  (II)  Jy.    1890,  T.  J.  McElligott  ;   Hennepin,  (III) 

-  1890,  E.  A.  Cuzner  ;  Traverse,  (III)  S.  1893,  Shel- 

don 7366;  Ramsey,  (III)  S.  1898,  Freeman  ;  --  ,  (III) 

1893,  Sheldon,  5823,  6175  and  (II,  III)  7195  ;  Hennepin, 

(II,  III)  O.  1889,  MacMillan. 

Helianthus   sp.   indet.  :    Traverse,   (III)  S.  1893,  Sheldon 

7378;  -  >  (HI)  l893>  Sheldon  7136  and  6067. 
10.  P.  Chondrillse  CORDA,  Icon.  Fung.  4:  15.     1840. 

The  Minnesota  specimens  of  the  Puccinia  on  Lactuca  ex- 
hibit morphological  characters  which  according  to  Jacky  (1.  c.) 
distinguish  this  form  from  those  on  Prenanthes.  The  teleuto- 
spore  has  no  well-developed  papilla  and  no  constriction  and  is 
elliptical  in  form.  The  germ  pores  are  irregular  in  position, 
often  occurring  at  the  summit. 

On  Lactuca  -pulchella  (PURSH)  DC.  :  -  ,  (III)  S.  1893, 

Sheldon. 

Lactuca  sp.  indet.  :  Traverse,  (III)  S.  1893,  Sheldon  7241 
and  7125. 

B.   HETER-EUPUCCINIA. 

n.  P.  angustata  PECK,  Rep.  N.  Y.  St.  Mus.  Nat.  Hist.  25  :  123. 


On  Sctrpus  atrovirens  MUHL.  :  Ramsey,  (II,  III)  O.  1898, 

Freeman. 
Lycofais  virginicus  L.  :   St.   Louis,  (I)  Jy.   1886,   Holway 

216;  Blue  Earth,  (I)  Je.  1891,  Sheldon  484. 
This  species  is  connected  with  ^Ecidium  lycopt  Ger.  accord- 
ing to  Arthur  (1.  c.). 

12.  P.  caricis  (SCHUM.)  REBENT.  Prod.  Fl.  Neom.  356.     1804. 

The  connection  of  ^Ecidium  urticce  Schwein.  on  Urtica  with 

P.  caricis  has  been  confirmed  for  American  specimens.     (Ar- 

thur, 1.  c.)     The  aecidia  are  found  on  both  lamina  and  petiole. 


Freeman:    MINNESOTA  UREDINE^E.  547 

On  Carcx  castanca  WAHL.  :  Lake,  (III)  Je.  1893,  Shel- 
don 4822. 

Carex  utriculata  BOOTT  :  Houston,  (III)  My.  1900, 
Lyon  537. 

Carex  sp.  indet.  :  — ,  (III)  1893,  Sheldon  7135,  7368 
and  7122;  Hennepin,  (III)  O.  1898,  Freeman;  Henne- 
pin,  (III)  S.  1900,  Freeman  786^. 

Urtica  gracilis  AIT.  :  Ramsey,  (I)  My.  1899,  Freeman 
322;  Waseca,  (I)  Je.  1891,  Sheldon  206  and  Taylor 
302. 

Urtica  sp.  indet.  :   Hennepin,  (I)  1893,  Sheldon  5968  and 

5967- 

13.  P.  phragmitis  (SCHUM.)  KORN.  Hedw.  15:  179.     1876. 
In  Europe  it  has  been  demonstrated  by  several  investigators 

that  the  aecidium  on  various  species  of  Rumex  belongs  to  P. 
phragm i/is.  According  to  the  recent  culture  experiments  of 
Arthur  (1.  c.)  aecidia  were  easily  obtained  upon  Rumex  crtspus 
and  Rumex  obtusifolius.  According  to  the  same  author  no  au- 
thentic and  undoubted  record  of  the  collection  of  ^Ecidntm 
rubcllum  GMEL.  has  been  reported  upon  American  species  of 
Rumex.  A  Minnesota  specimen  of  Riimex  britannica  L.  was 
collected  in  1899  (Freeman  471)  upon  which  several  groups  of 
aecidia  were  found.  Upon  comparison  with  sEcidmm  rubellum 
on  R.  hydrolapathum  *  the  two  specimens  were  found  to  agree 
perfectly  in  all  morphological  characters. 

The  spots  are  circular  and  usually  of  a  reddish  color,  not 
swollen.  Pseudoperidia  on  lower  surface  of  the  leaf  somewhat 
crowded,  leaving  usually  a  free  central  area,  flat,  cup-shaped  with 
revolute  lacerate  margin.  Spores  17— 23  //  x  n— 17 /^.  It  seems 
very  probable  therefore  that  this  ascidium  on  Rumex  britannica 
belongs  to  P.  phragmitis.  Arthur's  explanation  that  the  aecid- 
ium  on  Rumex  has  up  to  this  time  been  overlooked  is  therefore 
probably  correct. 

On  Rumex  britannica  L.  :  Ramsey,  (I)  Je.  1899,  Freeman 
471. 

Phraomites -phragmites  (L.)  KARST  :  ?(IH)  T893,  Shel- 
don 7119. 

14.  P.  rhamni  (P.)  WETTST.  Verh.  Zool.-Bot.  Ges.  Wien.  35: 

545.     1885.     (P.  coronata  CDA.) 

*  Krieger  Fung.  Sax.  no.  853. 


548  MINNESOTA    BOTANICAL    STUDIES. 

On  Avena  sativa  L.  :  Brown,  (II)  Jy.  1891,  Sheldon  1045. 

Rhamnus  alnifolia  L'HER.  :  Houston,  (I)  Ag.  1900,  Lyon 

546  ;  Ramsey,  (I)  My.  1899,  Freeman  316  ; ,  (I)  1893, 

Sheldon  5963  ;  Hennepin,  (I)  My.  1891,  Sheldon  5969. 

15.  P.    poculiformis  QACQ^)  WETT.    Verb.   Zool.-Bot.   Ges. 

Wien.  35:  544.      1885.     (P.  graminis  PERS.) 
There  is  no  good  reason  for  not  accepting  Wettstein's  name. 
The  species  is  exceedingly  abundant  although  but  few  collec- 
tions have  been  made. 

On  Avena  sativa  L.  :  Brown,  (III)  Jy.  1891,  Sheldon  1104; 
Goodhue,  (II)  Ag.  1893,  Anderson  710. 

Undetermined  grasses  : ,  (II,  III)  1893,  Sheldon  7120  ; 

,  (II),  1893,  Sheldon  7126. 

16.  P.  rubigo-vera  (DC.)  WINT.  Die  Pilze  i1 :   217.     1884. 
On    Triticum  vulgare  L.  :   Kandiyohi,  (II)  Jy.  1892,  Frost 

285   and  (II,  III)  283^;  Goodhue,  (II,  III)  Ag.   1893, 

Anderson  711  ;  Traverse,  (III)  S.  1893,  Sheldon  7387. 

Hordeum  vulgare  L.  :  Waseca,  (II,  III)  Je.  1891,  Sheldon 

529- 

C.  BRACHYPUCCINIA. 

17.  P.  hieracii  (SCHUM.)  MART.  Flora  Mosq.  226.     1817. 
Jacky's  results  (1.  c.)  can  be  utilized  only  in  separating  out 

the  form  on    Taraxacum.     Those  on  Hieracium  and   Carduus 

require  further  cultural  investigation  on  American  specimens. 

On    Hieracium    canadense   MICHX.  :    Hennepin,    (III)  O. 

1889,  MacMillan. 
Carduus  sp.  indet.  :  Winona,  (II)  Ag.  1888,  Holzinger  41  ; 

,  (II,  III)  1893,  Sheldon  6097  and  6059;  Traverse, 

(II,  III)  S.    1893,   Sheldon   7398;   Ramsey,  (II,  III)  S. 
1898,  Freeman. 

18.  P.  taraxaci  PLOWRIGHT,  Brit.  Ured.  and  Ustil.  :   186  and 

187.     1889. 

This  species  seems  to  be  founded  on  negative  results  of 
Plowright  and  recently  of  Jacky.  Inoculations  with  Puccinias 
from  other  composites  gave  in  all  cases  negative  results. 

On  Taraxacum  taraxacum  (L.)  KARST.  :  Traverse,  (II)  S. 
1893,  Sheldon  7324;  Hennepin,  (II)  Ag.  1883,  Arthur; 
Ramsey,  (II)  My.  1899,  Freeman  360 ;  Hennepin,  (II, 
III)  S.  1898,  Freeman. 


Freeman:    MINNESOTA  UKEDINE^E.  549 

D.  PUCCINIOPSIS. 

19.  P.  grossulariae  (GM.)  WINT.  Die  Pilze  i1 :   198.     1884. 
On  Ribcs  rub-rum  L.  :   St.  Louis,  (III)  Jy.  1886,  Holway 

213. 

20.  P.  podophylli  SCHWEIN.  Syn.   Fung.  Car.   Sup.  :    72  No. 

489.     1822. 

On  Podophyllum  f  citatum  L.  :  Winona,  (I)  My.  1889, 
Holzinger;  Winona,  (III)  Je.  1889,  Holzinger ;  Hous- 
ton, Je.  1899,  Lyon  (I)  2  and  (III)  80. 

E.  HEMIPUCCINIA. 

21.  P.  polygoni-amphibii   P.  Syn.  Meth.  Fungi   227.     1801. 
Very  widely  distributed  and  abundant. 

On  Polygonum  amphibium  L.  :  Hennepin,  (II,  III)  S. 
1889,  MacMillan; ,  (II,  III)  1893,  Sheldon  6076. 

Polygonum  hartivrightii  A.  GRAY  :  Hennepin,  (II,  III) 
Ag.  1883,  Arthur;  Hennepin,  (II,  III)  S.  1889,  Mac- 
Millan;  Otter  Tail,  (II,  III)  Ag.  1892,  Sheldon  3897; 
Traverse,  (II,  III)  S.  1893,  Sheldon  7377. 

Polygonum  enter  sum  (MiCHX.)  BRITTON  :  Hennepin, 
(II,  III)  Ag.  1883,  Arthur;  St.  Louis,  (II)  Jy.  1886, 
Holway  50  and  (II,  III)  Holway  48  ;  Winona,"  (II,  III) 
S.  1888,  Holzinger  252. 

Polygonum  sp.  indet.  :  Lincoln,  (II,  III)  Ag.  1891,  Sheldon 
1549  and  T57°;  -  — »  (m)  l893»  Sheldon  7137. 

22.  P.  argentata  (SCHULTZ)  WINT.  Die  Pilze  i2:  194.     1884. 
On    Impaticns   biflora   WALD.  :    Ramsey,   (III)  S.    1898, 

Freeman. 

23.  P.  pruni-spinosse  PERS.  Syn.   Meth.  Fung.   226.     1801. 
On  Primus  -puniila   L.  :    Crow  Wing,  (II,  III)  Ag.  1890, 

MacMillan  and  Sheldon  92. 

24.  P.  kuhniae   SCHWEIN.  Syn.  Fung.   Am.  Bor.   296.     1834. 
On   Kuhnia    eupatorioides    L.  :    Ramsey,    (III)   S.    1898, 

Freeman. 

25.  P.   tomipara  TREL.  Trans.  Wise.  Acad.  Sci.  A.  and  L. 

6:  23.     1884. 

On  Bromus  ciliatus  L.  :  St.  Louis,  (II)  Jy.  1886,  Holway 
12.  Also  reported  at  Detroit  (Becker  Co.)  in  Ell.  and 
Ever.  N.  A.  Fungi  No.  1842.  1884. 


550  MINNESOTA    BOTANICAL    STUDIES. 

26.  P.  sorghi  SCHWEIN.  Syn.  Fung.  Am.  Bor.  295.    No.  2910. 

1831. 

On  Zea  mays  L.  :  St.  Louis,  (II)  Jy.  1886,  Holway  136; 
—  ,  (II)    1893,  Sheldon  6148;   Brown,  (II)  Jy.  1891, 

Sheldon  1065. 
Sorghum  sp.  indet.  :   Hennepin,   (III)  O.,    1890,   Sheldon 

5742- 

27.  P.  emaculata   SCHWEIN.  Syn.  Fung.  Am.   Bor.  295.  No. 

2912.     1834. 

On  Panicum  capillare  L.  :  Lincoln,  (III)  Ag.  1891, 
Sheldon  1530;  Traverse,  (III)  S.  1893,  Sheldon  7375 
7376. 

F.  MlCROPUCCINIA. 

28.  P.  thalictri  CHEV.  Fl.  Paris,  i  :  417.     1826. 

On  Thalictrum  sp.  indet.  :  Cass,  (III)  Jy.  1893,  Ander- 
son 573. 

G.  LEPTOPUCCINIA. 

29.  P.  asteris  DUBY,  Botan.  Gallic.  2  :  888.     1828. 

On  Aster  macrophyllus  L.  :  St.  Louis,  Jy.  1886,  Holway 

37  ;  Cass,  Jy.  1893,  Ballard  1652. 
Aster    sagittlfolius  WILLD.  :     Houston,  Ag.   1899,  Lyon 


Aster  sp.  indet.  :  Lake,  Jy.  1886,  Holway,  273. 
There  is  a  great  similarity  in  the  spots,  sori  and  spores  of 
P.  asteris  on  A.  macrophyllus  to  those  of  P.  xanthti  Schwein. 

30.  P.  circaeae  PERS.  Tent.  Disp.  Meth.  39.     1797. 

On  Circcea  alpina  L.  :   St.  Louis,  Jy.  1886,  Holway  214. 
Circaa  lutetiana  L.  :  Winona,  Jy.,   1888,  Holzinger  206 

and  132  ;  Pope,  Jy.  1891,  Taylor  887. 
Circaa  sp.  indet.  :  Cass,  Jy.  1893,  Ballard  1655. 

31.  P.  ornata  ARTH.  &  HOLW.  Rep.  Bot.  Wk.  in  Minn,  for  year 

1886,  3:  30.      1887. 
On  Rumex  britannica  L.  :  St.  Louis,  (III)  Jy.  1886,  Hol- 

way 223. 
Rumex  sp.  indet.  :   Crow  Wing,  (III)  Ag.  1890,  MacMillan 

and  Sheldon. 

H.  LIFE  HISTORIES  IMPERFECTLY  KNOWN. 

32.  P.  variolans  HARK.  Bull.  Calif.  Acad.  Sci.  i  :  35.     1884. 
On  Eriocarpum  spinulosum  (NuTT.)    GREENE  :  Traverse, 

(III)  S.  1893,  Sheldon  7101. 


Freeman :   MINNESOTA  UREDINE^-E.  551 

33.  P.  nardosmiae  E.  &  E.  Journ.  Myc.  i :  85.     1885. 

On  Petasitcs  palmata  (Arr.)  A.  GRAY:  St.  Louis,  (III)  Jy. 
1886,  Holway  232. 

34.  P.    porphyrogenita   CURT.  Thiim.  Myc.    Univ.  No.    545. 

1876. 

On  Corntis  canadensis  L.  :   St.  Louis,  (III)  Jy.    1886,  Hol- 
way 108;  St.  Louis,  (III)  Je.  1893,  Sheldon  4622. 

35.  P.  hydrophylli PECK  &  CLINT.  Rep.  N.  Y.   St.  Mus.  Nat. 

Hist.  30:  54.     1879. 
On    Hydr&pkyllwm   -virgin! cum   L.  :   Hennepin,  (III)  My. 

1891,  Sheldon    1968  and  1981  ;    Meeker,  (III)  Je.  1892, 
Frost  37  ;  Ramsey,  (III)  My.  1892,  Sheldon  1969  ;  Aitken, 
(III)  Je.  1892,  Sheldon  2072  ;  -    — ,  (III)  1893,  Sheldon 
5803;    Hennepin,   (III)  My.  1899,  Freeman  310. 

36.  P.  tiarellae  B.  &  C.  N.  A.  Fungi,  No.  549.     1881. 

On  Mitclhi  unda^L.  :  St.  Louis,  (Ill)Jy.  1886,  Holway  62. 

37.  P.  halenise  ARTH.  &  HOLWAY,  Rep.  Bot.  Wk.  in  Minn. 

3:  30.     1886. 

On  Tetragonanthus  deflcxus  (J.  E.  SMITH)  KUNTZE  :  St. 
Louis,  (III)  Jy.  1886,  Holway  100. 

38.  P.  mesomegala  B.  &  C.  Grev.  3:  53.     1874. 

On  Clintonia  borcalis  (AiT.)  RAF.  :  St.  Louis,  (III)  Jy. 
1886,  Holway  18. 

39.  P.  xanthii  SCHWEIN.  Syn.  Fung.  Car.  Sup.  500.     1822. 
Berkeley  and  Ravenel  have  described  a  variety  ambrosia. 

The  spores  on  Ambrosia  do  not,  however,  differ  morpholog- 
ically from  those  on  Xanthiwn* 

QftXanthium  canadense  MILL.  :  Hennepin,  (III)  O.  1889, 
MacMillan ;  Brown,  (III)  Jy.  1891,  Sheldon  861  and 
1008;  Carver,  (III)  Jy.  1891,  Ballard  756;  Lincoln, 
(III)  Ag.  1891,  Sheldon  1262;  Otter  Tail,  (III)  Ag. 

1892,  Sheldon  3898;  Traverse,  (III)  S.  1893,  Sheldon 
7259;  Hennepin,  (III)  S.  1898,  Butters;  Ramsey,  (III) 
Jy.  1899,  Freeman  531  ;   Houston,  (III)  Ag.  1899,  Lyon 

458. 

Ambrosia  trifida  L.  :  Lincoln,  (III)  Ag.  1891,  Sheldon 
1266;  Brown,  (III)  Ag.  1891,  Sheldon  1227. 

Gymnoconia  LAGERH. 

Separated  from  Puccinia  on  account  of  the  aecidia,  which 
are  destitute  of  a  pseudoperidial  wall  and  are  at  first  covered 
only  by  the  epidermis  of  the  host. 


552  MINNESOTA    BOTANICAL    STUDIES. 

i.  Gymnoconia  interstitialis  (SCHLECT.)  LAGERH.  Ured.  Herb. 
El.  Fr.  Tromso.  Mus.  Arshefter  17  :  84.     1894.     (Cceoma 
nitcns  SCHWEIN.  and  Puccinia  pecki ana  HOWE.) 
The  aecidial  stage  (Cceoma  nitens  Schwein.),  the    common 
raspberry  rust,  is  abundant  throughout  the  State.     The  teleuto- 
spores  have  not  yet  been  found  in  Minnesota. 

On  Rubus  strigosus  MICHX.  :    Mille  Lacs,  (I)   Jy.   1892, 

Sheldon  2709. 
Rubus    villosus    AIT.  :    Kanabec,    (I)   Jy.    1892,    Sheldon 

2908;  Mille  Lacs,  (I)  Jy.  1892,  Sheldon  2713. 
Rubus  canadensis  L.  :   Le  Sueur,  (I)  Je.  1891,  Sheldon  52  ; 
Aitkin,   (I)  Je.  1892,  Sheldon  2061  and  2145  ;  Pine,  (I) 
Je.  1899,  Freeman  499. 

Rubus  sp.  indet.  :  Winona,  (I)  Je.  1888,  Holzinger  235  ; 
Cass,  (I)  Je.  1893  ;  Ballard  1393  ;  Houston,  (I)  Je.  1899, 
Lyon. 

Uropyxis  SCHROETER. 

The  puccinia  on  Amorpha  species  (P.  amorpha  Curt.)  has 
here  been  retained  under  Schroeter's  genus  Uropyxis.  The 
differences  between  Uropyxis  and  typical  Puccinia  are  no  less 
than  those  between  Phragmopyxis  and  Phragmidium.  On  the 
other  hand  P.  dcglubens  might  be  included  among  the  Puccinice 
without  much  more  serious  objection  than  can  be  raised  against 
Puccinia  elymi  Westd.  Puccinia  petalostemonis  Farlow  has  a 
slight  gelatinous  exospore  which  certainly  indicates  relationship 
with  the  Puccinia  on  Amorpha.  Phragmopyxis  according  to 
Dietel  *  is  more  closely  related  to  Uropyxis  than  to  Phragmi- 
dium. I  have  seen  biseptate  spores  in  Puccinia  petalostemonis 
Farl.f  The  forms  on  the  Leguminosse,  therefore,  seem  to 
form  a  natural  group  with  sufficient  distinguishing  characters  to 
separate  them  generically  from  the  typical  Puccinia.  This  is 
Schroeter's  genus  Uropyxis.  Puccinia  petalostemonis  Farl. 
connects  it  with  the  true  Puccince. 

i.  U.  amorphae  (CURT.)  SCHROET.  Hedw.  15:   165.     1875. 

On  Amorpha  canescens  PURSH  :  Winona,  (III)  S.  1888, 
Holzinger;  Otter  Tail,  (III)  Ag.  1892,  Sheldon  3650; 
Traverse,  (III)  S.  1893,  Sheldon  7111  and  7180; 
Ramsey,  (III)  S.  1898,  Freeman  ;  Pope,  (III)  Ag.  1891,. 
Taylor  1182. 

*  1.  c.,  70. 

t Ellis.     N.  A.  Fungi,  No.  1844. 


Freeman:    MINNESOTA  UREDINE^E.  553 

Amorpha  fruticosa  L.  :  Traverse,  (III)   S.   1893,  Sheldon 
7243;   Brown,   (III)  Jy.   1891,   Sheldon  985;   Chisago, 
(III)   S.    1891,  Sheldon   4245;   Traverse,  (III)  S.  1893, 
Sheldon  7370. 
The  aecidial  and  uredo  forms  have  not  yet  been  collected. 

Gymnosporangium  DE  CANDOLLE. 

1.  G.  globosum  FARI..  Am.   Mem.   Bot.   Soc.  Nat.  Hist.  18. 

1880. 
Qnjunipcrusvirgimana  L.  :  Dakota,  (III)  Ap.  1900,  Lyon. 

2.  G.  nidus-avis  THAXTER,  Bull.   Conn.   Ag.  Ex.  Sta.   107: 

6.     1891. 

On   Junipcrus   virginiana    L.  :    Wright,   (III)   Je.    1899, 
Lyon  553  and  547  and  Freeman  697. 

3.  G.  clavariaeforme  (JACQ..)  REES.    Abh.  Naturf.  Gesell.  n  : 

21.     1869. 
On  branches  of  Juniperus  communis  L.  :  Hennepin,  (III) 

A.  1901,  Butters  97. 
Causes  fusiform  swelling  of  the  branches. 

4.  G.  juniperi-virginianae  SCHW.  Syn.  Fung.  Car.  Sup.  74. 

No.  504.      1822.     (G.  macropus  LK.) 
On  small  branches  of  Juniperus  virginiana  L.  :  Wright, 

(III)  A.  1901,  Freeman  978. 
Causes  swellings  known  as  "  cedar  apples." 

Phragmidium  LINK. 

1.  P.    potentillae   (P.)  KARST.  Fungi  Fen.  No.  94  and  593. 

(Hel.  Bid.)  Fen.  Nat.  o.  Folk.  19:   1871—23:  1873. 
On  Potentilla  pennsylvanica   strigosa   PURSH  :    Douglas, 
(III)  Ag.  1892,  Sheldon  3481. 

2.  P.  rubi-idaei  (P.)  KARST.  Helsing.  Bid.  Fin.  Nat.  o.  Folk 

19.     1871. 

On  Rubus  strtgosus  MICHX.  :  Aitken,  (II)  Je.  1892,  Shel- 
don 221 1  ;  St.  Louis,  (II,  III)  Jy.  1886,  Holway  205. 

Rubus  hispidus  L.  :  St.  Louis,  (II)  Jy.  1886,   Holway  20. 

Rubus  sp.  indet.  :  Winona,  (II)  S.   1888,  Holzinger. 

3.  P.  speciosum  FR.  Syst.  Myc.  3:  496.     1829. 

On  stems  of  Rosa  sp.  indet.  :  Wright,  (III)  O.  1896,  Wash- 
burn;  Ramsey,  (III)  S.  1899,  Wheeler. 

4.  P.    subcorticum   (SCHRANK.)    WINT.    Die  Pilze   i1 :     228. 

1884. 


554:  MINNESOTA    BOTANICAL    STUDIES. 

On  Rosa  acicularis  LINDL.  :  St.  Louis,  (II,  III)  Jy.  1886, 
Holway  187  and  247  ;  Ramsey,  (III)  My.  1899,  Free- 
man 350. 

Rosa  blanda  AIT.  :  Mille  Lacs,  (III)  Je.  1890,  Sheldon  2313. 

Rosa  sp.  indet.  :  Hennepin,  (III)  O.  1898,  Freeman ;  St. 
Louis,  (II)  Jy.  1886,  Holway  3  ;  Winona,  (II,  III)  S. 
1888,  Holzinger;  Hennepin,  (III)  O.  1898,  Freeman; 
Houston,  (III)  Je.  1899,  Lyon  26;  (III)  1893,  Sheldon 

7334- 

Triphragmium  LINK. 

i.  T.  clavellosum  BERK.  Gard.  Chron.     1857. 

On  Aralia  nudicaulis  L.  :  St.  Louis,  (III)  Jy.  1886,  Hol- 
way 17. 

JEcidium.     ISOLATED  FORMS. 

1.  A.  uvulariae  SCHWEIN.  Syn.  Fung.  Car.  Sup.  69,  no.  453. 

1822. 
On  Uvularia  -perfoliata  L.  :  Blue  Earth,  Je.  1891,  Sheldon 

295  ;  Chisago,  Je.  1892,  Taylor  1255. 
Uvularia  grandiftora  SM.  :  Ramsey,  Je.    1899,   Freeman 

403;  Pine,  Je.    1899,  Freeman  526;  Houston,  Je.  1899, 

Lyon  96;  Houston,  Je.  1900,  Lyon  552. 
Uvularia  sessilifolia  L.  :  Aitkin,  Je.  1892,  Sheldon  2066. 

2.  A.  iridis   GER.   Rep.   N.  Y.  St.  Mus.  Nat.  Hist.  25:  93. 

1870. 

On  Iris  versicolor  L.  :  Kanabec,  Jy.  1892,  Sheldon  2886  ; 
Houston,  My.  1900,  Lyon  534. 

3.  A.  convallariae  SCHUM.  Enum.  Plant.  Saell.  2:  224.     1803. 
On  Lilium  canadense  L.  :  Ramsey,  Je.  1899,  Freeman  400. 
Polygonatum    commutatum  (Sen.)  DIETR.  :   Ramsey,   Je. 

1899,  Freeman  404. 
Polygonatum  sp.  indet.  :  Houston,  Je.  1899,  Lyon  22. 

4.  A.  orobi  PERS.  Rom.  N.  Mag.  i  :  92.     1794- 
Probably  belongs  to  Uromyces  fabce  (P.)  DE  BARY. 

(?)On  Falcata  comosa  (L.)  KUNTZE  :  Pope,  Jy.   1891,  Mac- 

Millan  5. 
Apios  apios  (L.)  MAcM.  :  Brown,  Jy.  1891,  Sheldon  914. 

5.  A.  lupini  PECK,  Rep.  N.  Y.  St.  Mus.  Nat.  Hist.  46:  33. 

1893. 
On  Lupinus  -perennis  L.  :  Chisago,  Jy.  1892,  Taylor  1421. 


Freeman:    MINNESOTA  UREDINE^E.  555 

6.  A.  thalictri-flavi  (DC.)  WINT.  Die  Pilze  i1  :    269.     1884. 
On     Thalictrnin    dioicnm     L.:      Mille     Lacs,    Je.     1892, 

Sheldon  2306. 

Thalictrum  piirpnrasccns  L.  :  St.  Louis,  Jy.  1886,  Hoi- 
way  210  (A.  thalictri  Grev.)  ;  Ramsey,  Je.  1898,  Free- 
man 71  ;  Ramsey,  My.  1899,  Freeman  318. 

7.  A.  ranunculacearum  DC.  Fl.  Fr.  6:  97.     1815. 

On  Anemone  canadcnse  L.  :   Winona,  Je.  1889,  Holzinger, 

Ramsey,  Je.  1899,  Freeman. 
Anemone  quinquefolia  L.  :  Aitkin,  Je.  1891,  Sheldon  2312 

and  2108;  Mille  Lacs,  Je.  1892,  Sheldon  2566. 
Ranunculus  abortivus   L.  :   Wright,   My.    1899,    Freeman 

637^  ;  St.  Louis,  Jy.  1886,  Holway  212. 
The  aecidia  on   R,  abortivus  L.    are    not  diffused   over  the 
entire   surface  but  are   aggregated   on  definite  orbicular  spots 
which    are    at     first    reddish    but    become    yellow   in    drying. 
Spores  finely  tuberculate  polygono-spherical,  20-23  x  17  /*.     Not 
ranunculi  Schwein. 


8.  A.  punctatum  PERS.  Usteri  Ann.  Bot.  20:  135.     1796. 
On    Thalictrum  dioicum  L.  :  Chisago,  Je.    1899,  Freeman 

416. 

9.  A.  actaeae  OPIZ.  in  Wallr.  Fl.  Crypt.  Germ.  2:  252.    1833. 
On  Actcea  alba  (L.)  MILL.  :  Blue  Earth,  Je.  1891,  Sheldon 

392- 

10.  A.   cimicifugatum  SCHWEIN.   Syn.   Fung.  Am.  Bor.    293. 

No.  2876.     1831. 

On  Actaa  sp.  indet.  :  Pope,  Jy.  1891,  Taylor  889. 
Differs  from  A.  cimicifugatum  Schwein.  in  the  long  cylin- 
drical pseudoperidia  and  in  smooth  (or  almost)  spores.     Spores 
15-17  IJL  in  diameter. 

11.  A.  clematidis  DC.  Fl.  Fr.  2:  243.     1815. 

On  Clematis  virginiana  L.  :  Mille  Lacs,  Jy.  1892,  Shel- 
don 2764;  Brown,  Jy.  1891,  Sheldon  907;  Pope,  Jy. 
1892,  MacMillan  6  ;  Houston,  Je.  1899,  Lyon  106. 

12.  A.  fumariacearum  KELL.  &  SWINGLE,  Journ.  Myc.  4:  95. 

1888. 

On  Bicuculla  cucullaria  (L.)  MILLSP.  :  Aitkin,  Je.  1892, 
Sheldon  2203  ;  -  1893,  Sheldon  5959. 


556  MINNESOTA    BOTANICAL    STUDIES. 

13.  A.  geranii  DC.  Syn.  PI.  47.     1806. 

On     Geranium    maculatum    L.  :    Crow  Wing,    Je.    1892, 
Sheldon   2247  ;  Meeker,  Je.  1892,  Frost  61  ;  Houston, 
Je.    1899,  Lyon  95  ;  Ramsay,  My.  and  Je.  1899,  Free- 
man 329  and  419;  Winona,  Je.  1888,  Holzinger. 
Belongs  probably  to  Uromyces  geranii  (DC.)  Wint. 

14.  A.  impatientis  SCHWEIN.  Syn.  Fung.  Car.  674.  No.  442. 

1822. 

On  Impatiens  biflora  WALT.  :  Winona,  Je.  1889,  Hol- 
zinger;  Pope,  Jy.  1891,  Taylor  826;  Waseca,  Je.  1891, 
Sheldon  520;  Le  Sueur,  Je.  1891,  Sheldon  219;  Mille 
Lacs,  Je.  1892,  Sheldon  2495. 

Impatiens  sp.  indet.  :  Pope,  Jy.  1892,  MacMillan  3 ; 
Wright,  Je.  1900,  Freeman  686. 

15.  A.  verbenas  SPEG.  Fung.  Argent,  i :  56.     1880. 

On  Verbena  stricta  VENT.  :  Brown,  Jy.  1891,  Sheldon 
1080. 

16.  A.  jacobeae  GREV.  Fl.  Edin.  445.     1824.      (A.  senecionis 

DESMAZ.) 

In  European  specimens  connected  with  Puccinia  Schceleriana 
Plow,  et  Magn. 

On  leaves  of  Senecio  aureus  L.,  Ramsey,  My.  1899,  Free- 
man 328. 

17.  A.  compositarum  MART.  Fl.  Erlang.  314.     1817. 

The  aecidia  on  composites  vary  considerably  in  the  form  and 
color  of  the  spots,  the  arrangement  of  the  ascidia  and  in  the 
size  and  form  of  the  spores.  In  the  absence  of  necessary 
knowledge  of  the  life-histories  of  these  forms  only  a  temporary 
classification  is  possible.  Their  separation  into  varieties  based 
on  the  hosts  is  the  only  convenient  method. 

On    Hieracium    canadense    Michx.  :    Ramsey,   Je.    1899, 

Freeman  396;  Chisago,  Je.  1899,  Freeman  425. 
On  Adopogon  'virginicum  (L.)  KUNTZE  :  Ramsey,  Je.  1899, 

Freeman  392. 
Var.  erigerontis  WINT. 

On  Erigeron  annuus  (L.)  PERS.  :  Houston,  Je.  1899,  Lyon 

85- 
Var.  prenanthis  (P.)  WALLR.  Fl.   Crypt.  Germ.  no.  1773. 

1833- 

On  Nabalus  sp.  indet.  :  Houston,  Je.  1899,  Lyon  12  ;  Chi- 
sago, Je.  1899,  Freeman  423. 


Freeman :    MINNESOTA  UREDINE^E.  557 

Var.   lactucae  BURRILL,  Bull.  111.   St.  Lab.  Nat.   Hist.   2 : 

232.     1885. 
On  Lactuca  canadcnsis  L.  :  Pine,  Je.  1899,  Freeman  475  ; 

Ramsey,   Je.    1898,    Freeman  69 ;    Chisago,  Je.    1899, 

Freeman  418. 
Lactuca  ludoviciana  (NuTT.)  DC.  :   Le  Sueur,  Je.   1891, 

Sheldon  291^  and  244. 
Lactuca  sp.  indet.  :  Ramsey,  My.  Freeman  344 ;  Wright, 

My.  1900,  Freeman  594. 

Var.  liatrii  WEBBER,  Journ.  Myc.  5.     1889. 

On  Laciniaria  sp.  indet.  :  Pine,  Je.  1899,  Freeman  514. 

Var.  helianthi  BURRILL,  1.  c.  232. 

On  Hclianthus  divaricatus  L.  :  Blue  Earth,  Je.  1891,  Shel- 
don 281. 
Helianthus  sp.  indet.  :  Le  Sueur,  Je.  1891,  Taylor  265. 

Var.  eupatorii  (ScHw.)  BURRILL,  1.  c.  231. 

On  Eupatorium  -purpureum  L.  :  Ramsey,  Je.  1899,  Free- 
man 464. 

Eupatorium  perfoliatum  L.  :  Brown,  Ag.  1891,  Sheldon 
1058  ;  Waseca,  Je.  1891,  Sheldon  308  and  521  ;  Chisago, 
Je.  1892,  Taylor  1340. 

Eupatorium  ageratoides  L.  :  Ramsey,  Je.  1898,  Freeman 
70. 

18.  A.  asterum  SCHWEIN.  Syn.  Fung.  Car.  Sup.  67,  No.  444. 

1822. 
On   Solidago  flexicaulis  L.  :  Waseca,  Jy.    1891,    Sheldon 

657. 
Solidago  serotina  AIT.  :    Mille  Lacs,  Jy.   1892,  Sheldon 

2790. 
Solidago    sp.  indet.  :    St.   Louis,  Jy.   1886,    Holway    141 

(A.  compositaruiri] ;  Blue  Earth,  Je.  1891,  Sheldon  207  ; 

Mille  Lacs,  Je.  1892,  Sheldon  2451  ;  Wright,  My.  1900, 

Freeman  595. 
Aster  sagittifolius  WILLD.  :  Blue  Earth,  Je.  1891,  Sheldon 

258.    ' 

Aster  sp.  indet.:  Waseca,  Je.  1891,  Sheldon  566;  Hous- 
ton, Je.  1899,  Lyon  14;  Ramsey,  Je.  1899,  Freeman 

398. 

Euthamia  graminifolia  (L.)  NUTT.  :  Chisago,  Je.  1899, 
Freeman  430;  Chisago,  Je.  1892,  Taylor  1339. 


558  MINNESOTA    BOTANICAL    STUDIES. 

19.  A.  pustulatum  CURT.,  Rep.  N.  Y.  St.  Mus.  Nat.   Hist. 

23 :  60.     1869. 

On  Comandra  umbcUata  (L.)  NUTT.  :  Winona,  Je.  1888, 
Holzinger ;  Wright,  Je.  1900,  Freeman  691;  Ramsey, 
Je.  1899,  Freeman  394 ;  Pine,  Je.  1899,  Freeman  490 ; 
Houston,  Je.  1899,  Lyon  90.  Very  abundant  throughout 
the  State. 

20.  A.  jamesianum  PECK,  Bot.  Gaz.  5  :  34.     1880. 

On  Asclepias  syriaca  L.  :  Brown,  Jy.  1891,  Sheldon  1078. 
Asclepias  tuberosa  L.  :  Brown,  Jy.  1891,  Sheldon  787. 
Acerates   viridiflora  (RAF.)  EATOX :    Houston,  Je.    1899, 
Lyon. 

21.  A.  lysimachiae  (SCHL.)  WALLR.    Fl.  Crypt.  Germ.  No. 

1770.     1833. 

On  Steironcma  ciliatum  (L.)  RAF.  :  Pine,  Je.  1899,  Free- 
man 503. 

22.  A.  grossulariae  PERS.   Syn.  Meth.  Fung.  (?).     1801.     (A. 
grossularice  SCHUM.  Enum.  Plant.  Saell.  2  :   223.      1803.) 

Abundant  everywhere. 

On  Ribes  gracile  MICHX.  :  Winona,  My.  1886,  Holzinger ; 
Brown,  Jy.  1891,  Sheldon  826^;  Ramsey,  My.  1899, 
Freeman  315. 

Ribes floridum  L'HER.  :  Le  Sueur,  Je.  1891,  Sheldon  233  ; 
Blue  Earth,  Je.  1891,  Sheldon  375  ;  Ramsey,  My.  1899, 
Freeman  326;  Wright,  Je.  1900,  Lyon  550;  Houston, 
Je.  1899,  Lyon. 

Ribes  cynosbati  L.  :  Aitkin,  Je.  1892,  Sheldon  2291  ; 
Wright,  My.  1900,  Freeman  660  ;  Pope,  Jy.  1891,  Taylor 
935;  Chisago,  Je.  1892,  Taylor  1307. 

Ribes  sp.  indet.  :  Blue  Earth,  Je.  1891,  Sheldon  375; 
Hennepin,  My.  1891,  Sheldon;  Meeker,  Je.  1892,  Frost 
33;  Lake,  Je.  1893,  Sheldon  4928;  Hennepin,  1893, 
Sheldon  5966 ;  Houston,  Je.  1900,  Lyon  551;  Wright, 
My.  1900,  Freeman  598;  Houston,  Je.  1899,  Lyon  13. 

23.  A.  hydnoideum  B.  &  C.  Grev.  3:  61.     1874. 

On  Dirca palustris  L.  :  Chisago,  S.  1891,  Sheldon,  1984^  ; 
Crow  Wing,  Jy.  1893,  Ballard  1496  and  1646;  Wright, 
My.  1900,  Freeman  579. 


Freeman:    MINNESOTA  UREDINE^E.  559 

24.  A.  hydrophylli  PECK,  Rep.  N.  Y.  St.  Mus.  Nat.  Hist.  26: 

78.     1874. 

On  HydrophyUwn  -cirginicnm  L.  :  Mille  Lacs,  Je.  1891, 
Sheldon  2475  and  2822  ;  Ramsay,  Je.  1899,  Freeman 
397  ;  Wright,  My.  1900,  Freeman  600. 

25.  A.  pammelii  TREL.  Trans.  Wis.  Acad.  Sc.A.andL.  6:  33. 

1884. 
On  Euphorbia  corollata  L.  :   Houston,  Jy.  1899,  Lyon. 

26.  A.  peckii  DETONI,  Syll.  Fung.  7:  790.     1888. 

On  Onagra  biennis  (L.)  SCOP.  :  Pope,  Jy.  1891,  Taylor 
865;  Waseca,Je.  1891,  Sheldon  510  ;  Waseca,  Je.  1892, 
Taylor  479;  Ramsey,  Jy.  1898,  Freeman  72;  Hennepin, 
My.  1899,  Freeman  337  ;  Pine,  Je.  1899,  Freeman  508 
and  498. 

27.  A.  phrymae  HALST.  Journ.  Myc.  2:  52.      1886. 

On  Phryma  leptostachya  L.  :  Waseca,  Je.  1891,  Sheldon 
564;  Brown,  Ag.  1891,  Sheldon  1000. 

28.  A.  fraxini  SCHVVEIN.  Syn.  Fung.  Car.  Sup.  66.  No.  430. 

1822. 
On  Fraxinus  Americana  L.  :  Lincoln,  Ag.  1891,  Sheldon 

1520;  Brown,  Jy.  1891,  Sheldon  1205. 
Fraxinus  sp.   indet.  :    Brown,  Jy.    1891,   Sheldon    1076; 

Kandiyohi,  Jy.  1892,  Frost  292. 

Peridermium  LEV. 

1.  P.  balsameum   PK.  Rep.  N.  Y.  St.  Mus.  Nat.  Hist.  27: 

104.     1875. 

On  Abies  balsamea  (L.)  MILL.  :  St.  Louis,  Jy.  1886,  Hoi- 
way  208. 

The  spores  of  this  specimen  are  uniformly  smaller  than  those 
described  by  Peck  and  agree  more  nearly  with  those  of  A. 
elatinnm.  No  distortion  of  the  branches  has  been  reported. 
The  spores  measure  14  x  17— 2O//. 

2.  P.  abietinum  (A.  &  S.)  THUM.  var.  decolorans  THUM. 

On  Picea  mariana  (MILL.)  B.S.P.  :  St.  Louis,  Jy.  1886, 

Holway  93. 

The  spores  agree  with  those  of  the  ascidia  of  Chrysomyxa 
ledi,  to  which  this  species  supposedly  belongs. 


560  MINNESOTA    BOTANICAL    STUDIES. 

Uredo.     ISOLATED  FORMS. 

1.  U.  polypodii  (P.)  DC.  Fl.  Fr.  6:  81.     1815. 

On  Cystopteris  fragilis  BERNH.  :  Houston,  Jy.  1899,  Lyon 
217. 

2.  U.  agrimoniae-eupatoriae  (DC.)  WINT.  Die   Pilze   i1 :  252. 

1884. 

On  Agrimonia  hirsuta  (MuHL.)  BICKNELL  :   St.  Louis,  Jy. 
1886,  Holway  209. 


VOL.  II. 


MINNESOTA  BOTANICAL  STUDIES. 


PART  V. 


PLATE  XXXII. 


XXX.     A  NEW  SPECIES    OF   ALARIA. 


DE  ALTON  SAUNDERS. 


During  the  summer  of  1896  while  investigating  some  physio- 
logical problems  in  the  Hopkins  Sea-side  laboratory,  the  writer 
collected  an  Alaria  which  did  not  seem  to  agree  with  any  of 
the  described  species.  No  specimens  of  the  Pacific  coast  Ala- 
rias  were  at  hand  for  comparison  and  the  writer  being  loath  to 
add  further  to  the  synonymy  of  this  variable  group  laid  the 
plant  aside  until  a  favorable  opportunity  for  study  should  pre- 
sent itself.  Recently  a  specimen  of  this  plant  with  several  other 
species  of  Alaskan  algas  was  submitted  to  Dr.  Kjellman  who 
pronounced  it  a  new  species,  related  to  his  A.  -prcelonga  and 
A.  angusta. 

Alaria  curtipes  nov.  sp.     (Plate  XXXIII.) 

Plant  of  medium  size,  one  to  three  or  more  meters  long,  dark 
olive  brown,  coriaceous ;  stipe  very  short  (1-4  cm.  long),  firm, 
robust,  black,  narrowed  below,  but  little  flattened  above ;  rachis 
short,  somewhat  compressed,  gradually  passing  into  the  midrib  ; 
blade  linear  or  narrowly  lanceolate,  1-3  dcm.  wide,  narrowed 
above ;  midrib  prominent,  1-2  cm.  broad,  projecting  equally  on 
both  surfaces  of  the  blade,  quadrangular  in  cross  section  ;  spor- 
ophylls  ovate,  lanceolate  or  elliptical,  obtusely  rounded  above, 
2-3  cm.  wide,  7-15  cm.  long,  16-40  or  more  borne  sedately  on 
a  distinct  stalk  5-10  mm.  long ;  fruiting  area  confined  to  the 
lower  half  of  the  sporophylls. 

Abundant  on  exposed  rocky  points  on  the  central  Californian 
coast,  Monterey  bay,  Carmel  bay,  and  Point  Sur. 

A.  curtipes  is  related  to  A.  prcelonga  *  Kjellm.,  and  A.  an- 
gusta f  Kjellm.  but  according  to  Dr.  Kjellman's  comparison 
"  Differs  from  A.  prcelonga  in  its  broader  midrib  and  its  shorter 


*  Kjellman.     Om  Beringshafv.  Algflora,  p.  38,  T.  4,  Figs.  1-4. 
f  Ibidem,  p.  38,   T.  3,  Figs.    1-4. 

561 


562  MINNESOTA    BOTANICAL    STUDIES. 

and  more  robust  stipe.     It  differs  from  A.  angusta  especially  in 
the  form  of  the  sporophylls." 

EXPLANATION  OF  PLATE  XXXIII. 

Figure  i.  Mature  plant  reduced  yz  ;   a,    b,    c,    d,    young   plants, 
showing  different  stages  of  development. 

Figure  2.  Cross  section  of  sporophylls,  X  400. 
Figure  3.   Section  of  midrib,  x   2. 


XXXI.   A   PRELIMINARY   LIST   OF    MINNESOTA 
XYLARIACE^:. 


F.  K.  BUTTERS. 


During  the  past  fifteen  years  numerous  collections  of  Min- 
nesota fungi  have  been  made  by  E.  W.  D.  Holway,  Esq.,  Dr. 
A.  P. 'Anderson,  Messrs.  E.  P.  Sheldon,  E.  M.  Freeman  and 
others.  The  list  given  below  comprises  records  of  all  the 
Xylariacese  which  have  been  collected  within  the  State  and 
deposited  in  the  herbarium  of  the  University  of  Minnesota.  In 
each  case  the  county  in  which  the  collection  was  made  is  cited, 
with  the  date  of  collection,  and  such  other  information  as  is 
deemed  of  special  value.  A  list  of  the  fungi  collected  by 
Holway  was  included  in  Professor  J.  C.  Arthur's  report  on 
botanical  work  in  Minnesota  for  the  year  1886,*  but  they  are 
included  in  the  present  list  for  the  sake  of  completeness.  Some 
of  Mr.  Sheldon's  specimens  as  well  as  the  fungi  included  in 
Professor  Arthur's  list  have  been  previously  determined,  but  in 
all  cases  the  fungi  reported  in  the  following  list  have  been  ex- 
amined personally  by  the  author  and  he  takes  the  sole  respon- 
sibility for  their  correct  determination.  In  cases  where  the 
determinations  as  given  in  Arthur's  list  have  been  altered  the 
name  as  it  appears  in  that  list  has  been  inserted  in  parentheses 
after  the  citation  of  the  collection. 

In  all  cases  in  which  the  nomenclature  employed  departs 
from  that  found  in  Ellis  and  Everhart's  North  American  Py- 
renomycetes,  the  name  employed  in  that  work  is  inserted  as  a 
synonym. 

It  is  to  be  noted  that  the  specimens  distributed  by  Ellis  f  as 
Hypoxylon  rubiginosum  (Pers.)  Fr.  are  certainly  of  a  different 
species  from  those  distributed  by  de  Thiimen  \  and  other  Euro- 

*  Geological  and  Natural  History  Survey  of  Minnesota,  Bulletin  No.  3, 
Oct.  i,  1887. 

t Ellis  &  Everhart.      North  American  Fungi,  1949,  Fungi  Columbiani,  1324. 
\Mycotheca  unt'versatis,  1071. 

563 


564  MINNESOTA    BOTANICAL    STUDIES. 

pean  authors.  De  Thiimen's  specimens  are  cited  by  Winter  * 
and  are  probably  authentic.  The  specimens  distributed  by 
Ellis  cannot  be  Hypoxylon  rubiginosum  (Pers.)  Fr.  They 
agree  with  the  fungus  upon  Magnolia  described  by  Berkeley  f 
as  Hypoxylon  epiphceum  B.  &  C.  (in  some  works  spelled 
epiphlceum}  and  the  species  has  been  so  cited  in  the  following  list. 

In  all,  nineteen  species  of  Xylariaceae  are  included  in  the  list 
given  below.  They  are  distributed  among  five  genera  as  fol- 
lows :  Nummularia,  3  ;  Ustulina,  i  ;  Hypoxylon,  12  ;  Daldinia, 
2  ;  Xylaria,  i. 

Owing  to  the  somewhat  desultory  manner  in  which  these  col- 
lections of  fungi  have  been  made,  many  species  which  doubtless 
occur  in  the  State  have  not  been  collected  as  yet,  while  some  of 
the  more  abundant  and  more  noticeable  species  have  been  col- 
lected many  times. 

1.  Nummularia   nummularia  (BULLIARD)   SCHROET.     Krypt. 
Fl.  von  Pilze  II.  459.      1897.     (N.   bulliardi  TUL.) 

Hennepin,   April    1890,   Sheldon    14,    on    Qiiercus; \ 

Sheldon  575 1%  ;  Wright,  May  1900,  Freeman  658  on 
^iiercus;  Hennepin,  September  1900,  Butters  75,  on  Acer; 
Hennepin,  October  1900,  Butters  50,  on  Acer. 

2.  Nummularia   repanda   (FRIES)  NITSCHKE,  Pyrenomycetes 
Germanici,  p.  57.      1867. 

Hennepin,  May  1893,  Sheldon  5428,  on   Cornusf  erumpent 
through  the  bark;  ,    Sheldon   5765;    ,   Sheldon 


;.  Nummularia  lateritia  ELLIS  &  EVERHART,  New  Species 
of  North  American  Fungi,  Proc.  Ac.  Nat.  Sci.  Philadel- 
phia, p.  144.  1893. 

On  bark. 

Hennepin,  May  1891,  Sheldon  4193,  on  Populus;  Hennepin, 
May  1891,  Sheldon  4197  ;  Ramsey,  May  1893,  Sheldon 
4327,  on  Acer ;  ,  Sheldon  5928,  on  Populus. 

\.  Ustulina  maxima  (HALLER)  SCHROTER,  Kryptogamen 
Flora  von  Schlesien,  Pilze  II.  p.  465.  1897.  (Ustulina 
vulgaris  TUL.) 


*Die  Pilze,  II.,  p.  860. 

t  Notices  of  North  American  Fungi,  Grevillea,  IV.  52.     1875. 
JMr.   Sheldon's  last  field  note-book  is  missing,  his  collections  concerning 
which  no  field  notes  can  be  found  are  indicated  as  above. 


Butters:    LIST  OF  MINNESOTA  XYLARIACE^E.  565 

Crow  Wing,  June  1892,  Sheldon  2238;  Ramsey,  August 
1893,  Sheldon  5528  ;  Hennepin,  June  1890,  Sheldon  5696  ; 

,  Sheldon  6138^3  ;  Hennepin,  May  1899,  Freeman 

306 ;  Ramsey,  June  1899,  Freeman  380 ;  Wright,  May 
1900,  Freeman  650^. 

Conidial  stage  :  Wright,  June  1900,  Freeman  684. 

5.  Hypoxylon  petersii  BERKELEY  &  CURTIS,  On  a  Collection 
of  Fungi  from  Cuba,  Journ.  Linn.  Soc.,  X.,  p.  384.      1869. 

Houston,  August  1899,  Wheeler  476,  on  decayed  log  ;  Hen- 
nepin, September  1900,  Freeman  702,  on  Jfyiercus. 

6.  Hypoxylon  fuscum  (PERS.)  FRIES,  Summa  Veg.  Scand.  p. 
384.     1849. 

On  bark. 

St.   Louis,  July    1886,    Holway   119,   on  Alnus;  St.  Louis, 

July  1886,  Holway  151,  on  Alnus;  Hennepin,  May  1891, 

Sheldon  4195  ;  Lake,  June  1893,  Sheldon  4749,  on  Alnus; 

Dakota,   July    1893,    Sheldon    5372;   Hennepin,    October 

1900,  Butters  51,  on  Ostrya. 

7.  Hypoxylon  commutation  NITSCHKE,  Pyrenomycetes  Ger- 
manici,  p.  33.      1867. 

On  bark. 

St.  Louis,  July    1886,  Holway  144,  on  Alnus;  Lake,  June 

1893,   Sheldon   4484,  on   Betula;    Hennepin,  May  1891. 

Sheldon  5904,  on  Tilia. 

8.  Hypoxylon  granulosum  BULLAIRD,  Champ.  Fr.  176.     1791 
(H.  multiforme  FR.)  Summa  Veg.  Scand.  p.  384.      1849. 

St.  Louis,  July  1886,  Holway  248,  on  Alnus  (Hypoxylon 
commutatum  Holwayanum  Sacc.  Holway);  St.  Louis, July 
1886,  Holway  262,  on  Betula;  St.  Louis,  June  1893,  Shel- 
don 4669;  Ramsey,  July  1893,  Sheldon  5490,  on  wood; 

,   Sheldon   6012,   on   Betula;  erumpent  through   the 

bark. 

9.  Hypoxylon  morsel  BERKELEY  &  CURTIS,  Notices  of  North 
American  Fungi,  Grevillea,  IV.,  p.  51.      1875. 

Erumpent  through  the  bark. 

St.  Louis,  July  1886,  Holway  41,  on  Betula  {Hypoxylon 
transversum  Schw.  Holway) ;  St.  Louis,  July  1886,  Hol- 
way 99,  on  Alnus;  Le  Sueur,  June  1891,  Taylor,  364; 
Hennepin,  May  1891,  Sheldon;  Crow  Wing,  June  1892, 


566  MINNESOTA    BOTANICAL    STUDIES. 

Sheldon  2054,  on  JQiicrcus;  Hennepin,  April  1891,  Shel- 
don 4178,  on  £>itercus;  Dakota,  July  1893,  Sheldon  5201, 
on  Qiiercus;  Hennepin,  May  1893,  Sheldon  5426,  on 

JQttcrctis;   ,   Sheldon  6138;  ,   Sheldon  6244;    on 

Qiiercus;  Wright,  June  1900,  Lyon542,  on  Rhns  ;  Chisago, 
September  1900,  Lyon  &  Butters,  on  Qiicrcus. 

10.  Hypoxylon  annulatum  (Scuw.)  MONTAGNE,  Sylloge  Cryp- 

togramarum,  p.  213.      1856. 
Cass,  August  1893,  Anderson  674,  on  wood. 

u.  Hypoxylon  marginatum  BERKELEY,   On   a   Collection   of 
Fungi   from   Cuba.       Part  II.,  Journ.   Lin.   Soc.,   X.,    p. 
499.      1869. 
Dakota,  July  1893,  Sheldon  5194  on  bark. 

12.  Hypoxylon   rubiginosum    (PERS.)    FRIES,    Summa   Veg. 
Scand.,  p.  384.     1849. 

On  wood. 

St.  Louis,  July  1886,  Holway  193  {Hypoxylon  ferrugineum 

Fr.  Holway  &  Ellis),  Ramsey,  July  1893,  Sheldon  5484; 

Wright,  May  1900,  Freeman  630. 

13.  Hypoxylon    perforatum    (ScHw.)    FRIES,    Summa   Veg. 
Scand.,  p.  384.     1849. 

,  Sheldon  5904^  ;  -     — ,  Sheldon  6133. 

14.  Hypoxylon  epiphaeum  BERKELEY  &   CURTIS,  Notices  of 
North  American  Fungi,  Grevillea,  IV.,  p.  52.      1875. 

Hypoxylon  rubiginosum  Ellis  &  Everhart,  North  American 
Fungi,  No.  1949,  not  H.  rubiginosum  (Pers.)  Fries. 

Hypoxylon  epiphlceum  B.  &  C. 

On  wood. 

Le  Sueur,  June  1891,  Taylor  435;  Brown,  July  1891, 
Sheldon  1027,  on  Tilia ;  Ramsey,  July  1893,  Sheldon 
5503,  on  Acer?  (young  form). 

15.  Hypoxylon  atropurpureum  (FRIES)   FRIES,  Summa  Veg. 

Scand.,  p.  384.      1849. 
On  wood. 
Le    Sueur,   June    1891,    Sheldon    94;    Brown,  July     1891, 

Sheldon     1027^,     on     Tilia;     Ramsey,     August     1893, 

Sheldon  5669;    ,   Sheldon  5751,   on  ££ticrctis ; , 

Sheldon  6236. 


Butters:    LIST  OF  MINNESOTA  XYLARIACE^E.  567 

16.  Hypoxylon  serpens  (PERS.)  FRIES,  Summa  Veg,  Scand., 

p.   384.     1849. 
St.  Louis,  July  1886,  Holway  265,  on  Popitlns. 

17.  Daldinia  tuberosa  (Scop.)  Voss.  Myc.  Carn.  180.     1891. 
(D.  concentrica  (BOLT.)  C.  &  N.)  Schema  di  Classifica- 
zione  degli  Sferiacei  Italic!  aschigeri,  Comment.  Soc.  Crit- 
tog.  Ital.,  I.,  p.  198.     1863. 

St.  Louis,  July  1886,  Holway  256 ;  Ramsey,  September 
1889,  Sheldon,  18  ;  Ramsey,  May  1890,  Sheldon  4340 ; 
Hennepin,  April  1891,  Sheldon  4191;  Blue  Earth,  June 
1891,  Sheldon  408;  Crow  Wing,  June  1892,  Sheldon 
2186;  Hennepin,  August  1893,  Sheldon  5590;  Henne- 
pin, September  1893,  Sheldon  5695  ;  Cass,  July  1893, 
Anderson  521  ;  Cass,  September  1898,  MacMillan  &  Free- 
man 108 ;  Wright,  May  1900,  Freeman  650 ;  Wright, 
May  1900,  Freeman,  654;  Chisago,  September  1900, 
Butters  85  ;  Hennepin,  September  1900,  Butters  71  ;  Hen- 
nepin, September  1900,  Butters  73. 

Conidial  stage. 

Waseca,  July  1891,  Taylor  670;  Chisago,  September  1900, 
Butters  87  ;  Hennepin,  October  1900,  Butters  62. 

18.  Daldinia    vernicosa    (Scnw.)    CESATI    &   DE    NOTARIS, 
Schema  di  classificazione  degli  Sferiacei  Italici  aschigeri, 
Comment.  Soc.  Crittog.  Ital.  I,  p.  198.     1863. 

Ramsey,  May  1890,  Sheldon;  Chisago,  September  1900, 
Butters  86;  Hennepin,  October  1900,  Butters  55  ;  Henne- 
pin, October  1900,  Butters  88. 

19.  Xylaria  clavata  (Scop.)  SCHRANCK,  Baierische  Flora,  II., 
p.   566.       1789.     ( Xylaria  -polymorpha  (Pers.)  Greville.) 

Hennepin,  April  1891,  Sheldon  4170;  Hennepin,  April  1891, 
Sheldon  4182  ;  Hennepin,  August  1893,  Sheldon  5581  ; 

,   Sheldon    6138^;    Hennepin,  July    1899,   Butters; 

Hennepin,  September  1900,  Freeman  784. 


VOL.  II. 


MINNESOTA  BOTANICAL  STUDIES. 


PART 


PLATE  XXXIII. 


XXXII.     A   CONTRIBUTION   TO  THE  KNOWLEDGE 

OF  THE  FLORA  OF   THE  RED  RIVER  VALLEY 

IN   MINNESOTA. 


W.  A.  WHEELER. 


During  August,  1900,  collections  were  made  by  the  Botanical 
Survey  in  the  valley  of  the  Red  River  of  the  North.  Professor 
Conway  MacMillan,  A.  S.  Skinner  andC.  J.  Hibbard  explored 
the  region  around  Crookston  through  Polk  and  Red  Lake 
counties  and  Professor  C.  A.  Ballard  visited  Kittson,  Marshall 
and  Otter  Tail  counties.  Professor  MacMillan's  party  visited 
Crookston,  Shirley,  Holmes  station  and  the  region  around 
Maple  lake  near  Dugdale  and  Mentor  in  Polk  county  and 
Thief  River  Falls,  Red  lake  Falls,  St.  Hilaire,  Wylie  and  Ives 
in  Red  Lake  county.  Professor  Ballard  visited  St.  Vincent, 
Humboldt,  Northcote,  Hallock  and  Kennedy  in  Kittson  county, 
Marshall  in  Warren  county  and  Fergus  Falls  in  Otter  Tail  county. 

The  collections  made  at  these  stations,  scattered  as  they  are 
through  the  Minnesota  part  of  the  Red  River  valley  give  a  good 
representation  of  the  late  summer  flora  of  this  region. 

The  following  plants  were  gathered  of  which  there  have  been 
no  definite  authentic  collections  previously  reported  from 
Minnesota. 

Puccincllia  airoides  (NuTT.)  WATS.  &  COULT. 

Elynius  macouni  VASEY. 

Scirpus  campestris  BRITTON. 

Juncus  dud  ley  i  WIEGAND. 

Rumex  occidentalis  S.  WATS. 

Chenopodinm  ambrosioides  L. 

Atriplcx  patula  L. 

Potentilla  ejfusa  DOUGL. 

Chamcerhodos  erecta  (L.)  BUNGE. 

Lappula  americana  (A.  GRAY)  RYDBERG. 

Chrysopsis  hispida  (HooK.)  NUTT. 

569 


570  MINNESOTA    BOTANICAL    STUDIES. 

As  a  result  of  the  reconnoissance  325  species  of  flowering 
plants  are  reported  below,  73  of  which  are  monocotyledons  and 

252  dicotyledons. 

Concerning  the  explorations  made  by  Professor  C.  A.  Ballard 
he  wrote  as  follows  under  date  of  September  i,  1900 : 

The  object  of  the  present  work  was  to  examine  certain  portions  of 
the  Red  River  valley  to  determine  (i)  whether  the  soil  of  the  sections 
under  examination  contained  saline  or  alkaline  ingredients  in  sufficient 
quantities  to  produce  a  distinctively  characteristic  vegetation,  and  (2) 
to  note  the  extent  of  territory  affected  by  such  conditions. 

I  found  it  very  difficult  to  obtain  definite  information  as  to  the  most 
pronounced  alkaline  regions  of  the  valley,  so  that  the  territory  covered 
is  a  part  only  of  the  sections  under  discussion. 

I  examined  first,  the  vegetation,  littoral  and  aquatic,  of  Mineral, 
Alkali  and  Horse  Shoe  lakes.  These  lakes  are  situated  south  and 
east  of  Fergus  Falls,  Otter  Tail  Co.,  and  are  more  or  less  strongly 
alkaline.  Of  the  three  examined  Mineral  lake  alone  has  a  vegetation 
differing  from  that  of  the  surrounding  country.  I  have  collected  in 
this  lake  Ruppia  occidental,  in  former  years  although  unable 
find  it  at  this  time.  It  is  without  doubt  growing  in  the  lake.  Around 
the  margin  of  the  lake  three  or  four  chenopods  grow  luxuriantly, 
next  spent  about  two  weeks  in  the  northern  part  of  the  valley  collect- 
ing in  the  vicinity  of  the  following  towns :  St.  Vincent,  Humboldt, 
Northcote,  Hallock  and  Kennedy  in  Kittson  Co.,  and  Warren  i 

Marshall  Co. 

The  monotonous  dead  level  of  the  prairie  is  broken  occasionally  by 
small  streams  each  with  its  fringe  of  trees.     The  surface  wells  of 
region  visited  are  alkaline,  some  of  them  decidedly  so.    This  shows  I 
entire  soil  to  be  alkaline  to  a  certain  extent.     These  wells  are  from  15 
to  25   feet  deep.     Many  deep   wells  have  been   sunk  throughou 
valley,  those  in  Kittson  Co.  generally  yielding  a  strong  brine  ( 
if  more  than  70  feet  deep.     I  visited  one  such  flowing  well   at  . 
boldt,  the  waters  of  which  had  killed  all  the  vegetation  for  rods  a 
the  path  of  the  flow.     These  conditions  are  so  recent  however  that  no 
marked  halophytic  vegetation  has  developed.     Near  a  similar  well  al 
Northcote  I  found  Salicornia  growing  abundantly  within  narrow  h 

Besides  these  localities  of  artificial  conditions  there  is  an  occasi. 
salt  spring  in  the  valley,  notably  one  on   "Two  Rivers"  some 
west  of  Hallock.     I  was  unable  to  reach  this  spring, 
depressions  in  the  surface  of  the  prairie  also  often  show  slight  i 
tions  of  an  alkaline  salt. 

At  Hallock  I  had  the  good  fortune  to  examine  the  herbarium  of 
G.  A.  Gunnarson,  the  Auditor  of  Kittson  Co.  This  herbarium  o: 
to  300  plants  represents  the  collections  of  several  years  in  the  imme- 


Wheeler:    THE  FLORA  OF  THE  RED  RIVER  VALLEY.         571 

diate  vicinity  of  Hallock.  These  plants  were  the  ordinary  types  of  the 
prairie  and  woodland  of  that  region.  One  plant,  however,  Plantago 
eriopoda,  is  worthy  of  note  as  being  a  salt  lover;  it  was  collected  15 
miles  southeast  of  Hallock  on  sandy  alkali  soil.  I  afterwards  found 
this  plant  growing  sparsely  six  miles  east  of  Warren. 

As  a  result  of  my  observations  I  draw  the  following  conclusions: 
The  soil  of  the  entire  valley  is  alkaline.  The  alkalinity  is  strongly 
marked  in  small  localities  only,  which  are  popularly  called  "  alkali 
spots."  In  Kittson  and  Marshall  counties  there  are  a  few  similar 
"  salt  spots,"  natural  and  artificial. 

There  are  few  halophytes  in  the  valley;  I  found  but  two  which  I 
considered  purely  halophytic,  these  are  B.  2680  Salicornia,  growing 
in  a  coulee  which  had  formerly  drained  a  salt  well  at  Northcote;  and 
B.  2789  Plantago,  growing  in  somewhat  elevated  sandy  soil  near 
Warren.  There  are  also  four  chenopods,  B.  2701,  B.  2702,  B.  2787, 
and  B.  2576,  which  are  semi-halophytic  in  character.  These  grow 
quite  generally  in  many  places  in  the  valley,  but  are  most  numerous 
and  luxuriant  on  the  beaches  of  alkali  lakes  and  ponds  and  in  the 
vicinity  of  alkali  spots. 

A  more  thorough  study  of  the  region  visited  will  doubtless  add  to 
the  list  of  halophytes  of  the  State  and  certainly  extend  the  range  of 
those  already  noted. 

The  principal  object  of  Professor  MacMillan,  in  his  visit  to 
the  valley,  was  to  secure  a  series  of  characteristic  photographs 
of  vegetation  and  portraits  of  plants  to  illustrate  the  flora  of  the 
region.  There  are  presented  herewith  some  views  selected  by 
him  and  made  under  his  direction  by  Mr.  C.  J.  Hibbard,  Pho- 
tographer of  the  Survey.  They  will  serve  to  give  an  idea  of  the 
vegetation-sheet  in  the  district  covered  by  the  list  and  will  indi- 
cate some  details  of  ecological  distribution  as  suggested  in  the 
descriptions  of  the  plates  written  by  Professor  MacMillan. 


LIST  OF  SPECIES. 

SPARGANIACE.E. 
Sparganium  eurycarpum    ENGELM.    in   A.   Gray,   Man.    Ed. 

2,  430.       1856. 

Coll.:     MacMillan    &    Skinner    131,    Maple    lake;    398, 
Holmes. 

Sparganium  simplex  HUDS.  Fl.  Angl.  Ed.  2,  401.     1788. 
Coll.:    Ballard  2581,  Humboldt. 


572  MINNESOTA    BOTANICAL    STUDIES. 

NAIADACEvE. 

Potamogeton  perfoliatus  L.  Sp.  PI.  126.     1753. 

Coll.:  Ballard  2651,  St.  Vincent;   MacMillan  &  Skinner 
173,  Maple  lake. 

Potamogeton  pectinatus  L.  Sp.  PL  126.     1753. 

Coll.:  Ballard  2652,   St.  Vincent;    MacMillan  &  Skinner 
172,  Maple  lake. 

SCHEUCHZERIACE^:.       . 

Triglochin  maritima  L.  Sp.  PI.  339.     1753. 

Coll.  :  MacMillan  &  Skinner  112,  Dugdale. 

ALISMACE^E. 

Alisma  plantago-aquatica  L.  Sp.  PI.  342.     1753. 

Coll.:    Ballard    2583,    Humboldt ;    2746,    Hallock ;    Mac- 
Millan &  Skinner  24,  417,  Crookston. 

Sagittaria  latifolia  WILLD.  Sp.  PI.  409.     1806. 

Coll.  :   MacMillan  &  Skinner  10,  Crookston  ;  370,  Holmes. 

Sagittaria  arifolia  NUTT.  ;  J.  G.  Smith,  Ann.   Rep.  Mo.  Bot. 

Gard.  6:  32.     1894. 

Coll.  :   Ballard  2580,  Humboldt;  2657,  St.  Vincent. 
Previously  reported  from  Minnesota  by  J.  G.  Smith  in  Ann. 
Rep.  Mo.  Bot.  Gard.  6:  33.      1895. 

NYMPKEACE.E. 

Nymphaea  advena  SOLAND,  in  Ait.  Hort.  Kew.  2  :  226.     1789. 
Coll.  :  MacMillan  &  Skinner  396,  Holmes. 

VALLISNERIACE^. 
Philotria  canadensis  (Micnx.)  BRITTON,    Science   (II)  2:5. 

1895. 

Coll.  :  MacMillan  &  Skinner  23,  Crookston. 

GRAMINE^. 

Andropogon  scoparius  MICHX.  Fl.  Bor.  Am.  i :  57.     1803. 
Coll.  :  MacMillan  &  Skinner  254,  Crookston. 

Andropogon  furcatus  MUHL.  :  Willd.  Sp.  PI.  4 :  919.     1806. 
Coll.:    Ballard    2598,    Humboldt;    2769,  Hallock;    Mac- 
Millan &  Skinner  357,  Shirley. 


Wheeler:   THE  FLORA  OF  THE  RED  RIVER  VALLEY.         573 

Chrysopogon  avenaceus  (MicHx.)  BENTH.  Journ.  Linn.  Soc.  19  : 

73.     1881. 
Coll.  :  MacMillan  &  Skinner  64,  Crookston. 

Panicum  crus-galli  L.  Sp.  PL  56.     1753. 

Coll.  :   Ballard  2589,  Humboldt ;   2699,  Northcote  ;  Mac- 
Millan &  Skinner  128,  129,  Maple  lake. 

Panicum  pubescens  LAM.  Encycl.  4:  748.     1797. 
Coll.  :  MacMillan  &  Skinner  138,  Maple  lake. 

Panicum  virgatum  L.  Sp.  PI.  59.     1753. 

Coll.:  MacMillan  &  Skinner  341,  342,  Crookston. 

Panicum  capillare  L.  Sp.  PI.  58.     1753. 

Coll.  :  Ballard  2541,  Fergus  Falls;  2663,  St.  Vincent. 

Chaetochloa  viridis  (L.)  SCRIBN.  U.  S.  Dept.  Agr.,  Div.  Agros. 

Bui.  4:  39.     1897. 
Coll.  :  Ballard  2525,  Fergus  Falls;  2686,  Northcote. 

Zizania  aquatica  L.  Sp.  PI.  991.     1753. 

Coll.  :   MacMillan  &  Skinner  134,  Maple  lake. 

Muhlenbergia  racemosa  (Micnx.)  B.S.P.  Prel.  Cat.  N.  Y.  67. 

1888. 

Coll.  :   Ballard  2693,  Northcote;   2754,  Hallock ;  MacMil- 
lan &  Skinner  99,  101,  Dugdale. 

Alopecurus  geniculatus  L.  Sp.  PI.  60.     1753. 

Coll.:     Ballard  2584,   Humboldt;    2747,   Hallock;    Mac- 
Millan &  Skinner  305,  306,  Crookston. 

Sporpbolus  brevifolius  (NuTT.)  SCRIBN.  Mem.  Torr.  Club,  5: 

39-     l895- 
Coll.  :  MacMillan  &  Skinner  329,  Crookston. 

Sporobolus  cuspidatus  (TORR.)  WOOD,  Bot.  &  Fl.  385.     1870. 
Coll.:     MacMillan    &    Skinner    275,    St.    Hilaire ;    384, 
Holmes. 

Sporobolus  heterolepis  A.  GRAY,  Man.  576.     1848. 
Coll.  :  MacMillan  &  Skinner  387,  Holmes. 

Agrostis  alba  L.  Sp.  PI.  63.     1753. 

Coll.  :  MacMillan  &  Skinner  233,  Crookston. 


574  MINNESOTA    BOTANICAL    STUDIES. 

Spartina  cynosuroides  (L.)  WILLD.  Enum.  80.     1809. 

Coll.:  Ballard  2585,  Humboldt;  2695,  Northcote ;  2762, 
Hallock;  2794,  Warren;  MacMillan  &  Skinner  52, 
Crookston. 

Bouteloua  oligostachya  (NuTT.)  TORR.  ;  A.  Gray,  Man.  Ed. 

2,  553-     1856. 
Coll.  :  MacMillan  &  Skinner  103,  Dugdale. 

Bouteloua  curtipendula  (Micnx.)  TORR.  Emory's   Rep.    153. 

1848. 
Coll.  :  MacMillan  &  Skinner  46,  Crookston. 

Beckmannia  erucaeformis  ( L. )  HOST,  Gram.  Austr.  3:5.    1805 . 
Coll.  :  Ballard  2588,  Humboldt ;   2634,  St.  Vincent ;   2714, 
Northcote;   2745,  Hallock;   2778,  Warren ;  MacMillan 
&  Skinner  100,  Dugdale;  361,  Crookston. 

Phragmites  phragmites  (L.)  KARST.  Deutsch  Fl.  379.    1880-83. 
Coll.  :  MacMillan  &  Skinner  394,  Holmes. 

Eragrostis  hypnoides  (LAM.)  B.S.P.  Prel.  Cat.  N.  Y.  69.    1888. 
Coll.  :  Ballard  2672,  St.  Vincent;  2751,  Hallock. 

Kceleria  cristata  (L.)  PERS.  Syn.  i :  97.     1805. 

Coll.:  MacMillan  &  Skinner  102,  Dugdale;  337,  Crook- 
ston. 

Panicularia  americana  (TORR.)  MAcM.  Met.  Minn.  Val.  81. 

1892. 
Coll.  :  Ballard  2582,  Humboldt. 

Puccinellia  airoides  (NUTT.)  WATS.   &  COULT.   in  A.  Gray, 

Man.  Ed.  6,  668.     1890. 
Coll.  :  Ballard  2528,  Fergus  Falls. 
Not  previously  reported  from  Minnesota. 

Bromus  ciliatus  L.  Sp.  PI.  76.     1753. 

Coll.:  Ballard  2782,  Warren;  MacMillan  &  Skinner  35, 
Crookston  ;  106,  Dugdale. 

Bromus  purgans  L.  Sp.  PL  76.     1753. 

Coll.:  Ballard  2666,  St.  Vincent;  2755,  Hallock ;  Mac- 
Millan &  Skinner  28,  Crookston. 

Bromus  kalmii  A.  GRAY,  Man.  600.     1848. 

Coll.  :  MacMillan  &  Skinner  105,  Dugdale. 

Agropyron  repens  (L.)  BEAUV.  Agrost.  146.     1812. 
Coll.  :  Ballard  2621,  Humboldt;  2715,  Northcote. 


Wheeler:    THE  FLORA  OF  THE  RED  RIVER  VALLEY.         575 

Agropyron  tenerum  VASEY,  Coult.  Bot.  Gaz.  10 :  258.     1885. 
Coll.  :  MacMillan  &  Skinner  304,  335,  Crookston  ;  Ballard 
2569,  St.  Vincent. 

Hordeum  jubatum  L.  Sp.  PI.  85.     1753. 
Coll.  :  Ballard  2520,  Fergus  Falls. 

Elymus  virginicus  L.  Sp.  PI.  84.     1753. 

Coll.  :  Ballard  2629,  St.  Vincent;  MacMillan  &  Skinner 
235,  Crookston. 

Elymus  canadensis  L.  Sp.  PI.  83.     1753. 

Coll.:  Ballard  2599,   Humboldt ;   2713,  Northcote ;  Mac- 
Millan &  Skinner  68,  Crookston  ;  267,  St.  Hilaire. 

Elymus  macouni  VASEY,  Bull.  Torr.  Club,  13:  119.     1886. 

Coll.  :  Ballard  2570,  St.  Vincent. 
Not  previously  reported  from  Minnesota. 

Hystrix  hystrix  (L.)  MILLSP.  Fl.  W.  Va.  474.     1892. 

Coll.:  MacMillan  &  Skinner   107,   Maple  lake;   265,  St. 
Hilaire. 

CYPERACEiE. 

Cyperus  diandrus  TORR.  Cat.  PL  N.  Y.  90.     1819. 
Coll.  :  Ballard  2537,  Fergus  Falls. 

Cyperus  speciosus  VAHL,  Enum.  2  :  364.     1806. 
Coll.  :  MacMillan  &  Skinner  130,  Maple  lake. 

Eleocharis  aciculails  (L.)  R.  &  S.  Syst.  2 :  154.     1817. 
Coll.  :  MacMillan  &  Skinner  141,  Maple  lake. 

Eleocharis  intermedia  (MUHL.)  SCHULTES,  Mant.  2  :  91.     1824. 
Coll.  :  Ballard  2656,  St.  Vincent. 

Scirpus  lacustris  L.  Sp.  PI.  48.     1753. 

Coll.  :  Ballard  2587,  St.  Vincent,  MacMillan  &  Skinner 
104,  Maple  lake. 

Scirpus  campestris  BRITTON,  in  Britton  &  Brown,  111.  Fl.  I : 

267.     1896. 

Coll.  :  Ballard  2539,  2544,  Fergus  Falls.     Not  previously 
reported  from  Minnesota. 

Scirpus  atrovirens  MUHL.  Gram.  43.     1817. 

Coll.:    MacMillan  &   Skinner   126,   127,  Dugdale ;    390, 
Holmes;  310,  Crookston. 


576  MINNESOTA    BOTANICAL    STUDIES. 

Carex  utriculata  BOOTT;  Hook.  Fl.  Bor.  Am.  2:  221.     1840. 
Coll.:    MacMillan   and  Skinner   139,   Maple  lake;    381, 
Holmes. 

Carex  retrorsa  SCHWEIN.  Ann.  Lye.  N.  Y.  i :  71.     1824. 
Coll.  :  MacMillan  &  Skinner  423,  Crookston. 

Carex  fusca  ALL.  Fl.  Fed.  2  :  269.     1785. 

Coll.  :  MacMillan  &  Skinner  133,  Maple  lake. 

Carex  cristatella  BRITTON,  in  Britton  &  Brown,  111.  Fl.  1 :  357. 

1896. 
Coll.  :  MacMillan  &  Skinner  249,  Red  Lake  Falls. 

Carex  sychnocephala  CAREY,  Am.  Journ.  Sci.  (II.)  4  :  24.    1847. 
Coll.  :  MacMillan  &  Skinner  307,  Crookston. 

ARACE^E. 

Arisaema  triphyllum  (L.)  TORR.  Fl.  N.  Y.  2  :  239.     1843. 
Coll.  :  MacMillan  &  Skinner  237,  Gentilly. 

Calla  palustris  L.  Sp.  PI.  968.     1753. 

Coll.  :  MacMillan  &  Skinner  174,  Maple  lake. 

Acorus  calamus  L.  Sp.  PI.  324.     1753. 

Coll.  :  MacMillan  &  Skinner  282,  Thief  River  Falls. 

LEMNACEJE. 
Lemna  trisulca  L.  Sp.  PI.  970.     1753. 

Coll.  :  Ballard   2654,  St-  Vincent ;  MacMillan  &  Skinner 
410,  Holmes. 

Lemna  minor  L.  Sp.  PI.  970.     1753. 
Coll.  :  Ballard  2546,  Fergus  Falls. 

JUNCACE^E. 

Juncus  balticus  WILLD.  Berlin  Mag.  3:  298.     1809. 

Coll.:    MacMillan    &    Skinner    136,    Maple    lake;    401, 
Holmes. 

Juncus  vaseyi  ENGELM.  Trans.  St.  Louis  Acad.   Sci.   2:450. 

1866. 
Coll.  :  MacMillan  &  Skinner  135,  Maple  lake. 

Juncus  dudleyi  WIEGAND,  Bull.  Torr.  Club,  27:  524.     1900. 
Coll.:    MacMillan   &     Skinner    137,    Maple    lake;    276, 
Wylie. 


Wheeler:   THE  FLORA  OF  THE  RED  RIVER  VALLEY.         577 

Not  previously  reported  from  Minnesota.  Many  specimens 
however  have  been  previously  collected  and  reported  as 
Juncus  tennis  Willd. 

Juncus  nodosus  L.  Sp.  PL  Ed.  2,  466.     1762. 

Coll.  :  Ballard  2741,  Hallock  ;  MacMillan  &  Skinner  373, 
Holmes. 

Juncus  torreyi  COVILLE,  Bull.  Torr.  Club,  22  :  303.     1895. 
Coll.  :   MacMillan  &  Skinner  340,  Crookston. 

Juncus  acuminatus  MICHX.  Fl.  Bor.  Am.  i :  192.     1802. 
Coll.  :   Ballard  2743,  Hallock. 

LILIACE^E. 

Allium  stellatum  KER,  Bot.  Mag.  pi.  1576.     1813. 

Coll.  :  MacMillan  &  Skinner  63,  Crookston  ;  87,  Dugdale. 

CONVALLARIACEJE. 

Vagnera  racemosa  (L.)  MORONG,  Mem.  Torr.  Club,  5:  114. 

1894. 
Coll.  :   MacMillan  &  Skinner  148,  Maple  lake. 

Vagnera  stellata  (L.)  MORONG,-  Mem.  Torr.  Club,  5  :  114.   1894. 
Coll.  :  MacMillan  &  Skinner  218,  Crookston. 

Unifolium  canadense  (DESF.)  GREENE,  Bull.  Torr.  Club,  15: 

287.     1888. 
Coll.  :  MacMillan  &  Skinner  147,  Maple  lake. 

Polygonatum  commutatum  (R.  &  S.)  DIETR.  ;  Otto  &  Dietr. 

Gartenz.  3  :   222.     1835. 
Coll.  :  MacMillan  &  Skinner  17,  Crookston. 

SMILACE^E. 

Smilax  herbacea  L.  Sp.  PI.  1030.     1753. 

Coll.  :  MacMillan  &  Skinner  18,  319,  Crookston;  Ballard 
2758,  Hallock. 

IRIDACE^E. 

Iris  versicolor  L.  Sp.  PI.  39.     1753. 

Coll.  :  MacMillan  &  Skinner  308,  Crookston. 

SALICACE^:. 

Populus  balsamifera  L.  Sp.  PI.  1034.     1753. 
Coll.  :  MacMillan  &  Skinner  278,  Ives. 


578  MINNESOTA    BOTANICAL    STUDIES. 

Salix  lucida  MUHL.  Neue  Schrift.  Ges.  Nat.  Fr.  Berlin  4  :  239. 

•pL  6.f.  j,     1803. 
Coll.  :  MacMillan  &  Skinner  366,  Holmes. 

BETULACE^E. 

Corylus  americana  WALT.  Fl.  Car.  236.     1788. 
Coll.  :  MacMillan  &  Skinner  415,  Crookston. 

Betula  papyrifera  MARSH.  Arb.  Am.  19.     1785. 
Coll.  :  MacMillan  &  Skinner  178,  Maple  lake. 

Betula  glandulosa  MICHX.  Fl.  Bor.  Am.  2:   180.     1803. 
Coll.  :  Ballard  2803,  Warren. 

Alnus  alnobetula  (EHRH.)  K.  KOCH,  Dendr.  2:  Part  i,  625. 

1872. 
Coll.  :  MacMillan  &  Skinner  151,  Maple  lake. 

FAGACE^E. 

Quercus  alba  L.  Sp.  PI.  996.     1753. 
Coll.  :  Ballard  2753,  Hallock. 

Quercus  macrocarpa  MICHX.  Hist.  Chen.  Am.  2.  pL  23.     1801. 
Coll.  :  MacMillan  &  Skinner  280,  Thief  River  Falls. 

HORACES. 

Humulus  lupulus  L.  Sp.  PI.  1028.     1753. 

Coll.  :  MacMillan  &  Skinner  26,  Crookston. 

URTICACE^E. 

Urtica  gracilis  AIT.  Hort.  Kew.  3:  341.     1789. 
Coll.  :  MacMillan  &  Skinner  199,  Crookston. 

Urticastrum  divaricatum  (L.)   KUNTZE,  Rev.  Gen.  PI.  635. 

1891. 
Coll.  :  MacMillan  &  Skinner  193,  Crookston. 

POLYGONACE.E. 

Rumex  verticillatus  L.  Sp.  PI.  334.     1753. 

Coll.  :   MacMillan  &  Skinner  158,  Maple  lake. 

Rumex  occidentalis   S.    WATS.    Proc.   Am.    Acad.    12 :   253. 

1876. 

Coll.  :  MacMillan  &  Skinner  270,  Thief  River  Falls. 
Not  previously  reported  from  Minnesota. 


Wheeler:   THE  FLORA  OF  THE  RED  RIVER  VALLEY.         579 

Rumex  crispus  L.  Sp.  PL  335.     1753. 

Coll.:  Ballard   2628,81.   Vincent;   2777,  Warren;    Mac- 
Millan  &  Skinner  189,  Crookston  ;  389,  Holmes. 

Rumex  persicarioides  L.  Sp.  PI.  335.     1753. 

Coll.:  Ballard  2638,81.    Vincent;  MacMillan  &  Skinner 
155,  Maple  lake. 

Polygonum  emersum  (Micnx.)  BRITTON,  Trans.  N.  Y.  Acad. 

Sci.  8:   73.      1889. 
Coll.  :  MacMillan  &  Skinner  412,  Crookston. 

Polygonum  lapathifolium  L.  Sp.  PL  360.     1753. 

Coll.  :  MacMillan  &  Skinner   2,  Crookston  ;   294,   Maple 
lake;  Ballard  2590,  Humboldt ;   2673,  St.  Vincent. 

Polygonum  persicaria  L.  Sp.  PL  361.     1753. 
Coll.  :  MacMillan  &  Skinner  6,  Crookston. 

Polygonum  punctatum  ELL.  Bot.  S.  C.  &  Ga.  i :  455.     1817. 
Coll.:    MacMillan  &  Skinner  293,    Maple  lake;  Ballard 
2530,  Fergus  Falls. 

Polygonum  littorale  LINK  in  Schrad.  Journ.  i :  54.     1799. 

Coll.  :  Ballard  2643,  2670,  St.  Vincent;   2685,  Northcote. 

Polygonum  erectum  L.  Sp.  PL  363.     1753. 
Coll.  :  Ballard  2720,  Kennedy. 

Polygonum  exsertum  SMALL,  Bull.  Torr.  Club,  21 :   172.     1894. 
Coll.  :   Ballard  2786,  Warren. 

Polygonum  ramosissimum  MICHX.  Fl.  Bor.  Am.  i :  237.  1803. 
Coll.  :  Ballard  2551,  2636,81.  Vincent;   2600,  Humboldt; 
MacMillan  &  Skinner  142,  Dugdale. 

Polygonum  convolvulus  L.  Sp.  PL  364.     1753. 

Coll.:  Ballard    2602,   Humboldt;    MacMillan   &  Skinner 
56,  Crookston. 

Polygonum  scandens  L.  Sp.  PL  364.     1753. 

Coll.  :  MacMillan  &  Skinner  167,  Maple  lake. 

CHENOPODIACE^E. 

Chenopodium  album  L.  Sp.  PL  219.     1753. 

Coll.:  Ballard  2552,  2575,  St.  Vincent;   2740,    Hallock ; 
2527,  Fergus  Falls  ;   2708,  Northcote  ;   2594,  Humboldt ; 
2721,  Kennedy;  MacMillan  &  Skinner  19,  Crookston. 
119,  Maple  lake. 


580  MINNESOTA    BOTANICAL    STUDIES. 

Chenopodium  glaucum  L.  Sp.  PL  220.     1753. 

Coll.  :  Ballard  2529,  2531,  Fergus  Falls  ;  2576,  St.  Vincent. 

Chenopodium    leptophyllum   (MooJ   NUTT.  :    Moq.    in    DC. 

Prodr.  13,  Part  2,  71.      As  synonym.      1849. 
Coll.  :  MacMillan  &  Skinner  332,  Crookston. 

Chenopodium  hybridum  L.  Sp.  PI.  219.     1753. 

Coll.:  Ballard  2558,  St.  Vincent;  MacMillan  &  Skinner 
225,  Crookston. 

Chenopodium  ambrosioides  L.  Sp.  PI.  219.     1753. 

Coll.  :  Ballard  2635,  St.  Vincent;  2595,  Humboldt ;   2687, 
Northcote;   2761,   Hallock ;   2787,  Warren;  MacMillan 
&  Skinner  424,  Crookston. 
No  previous  authentic  collection  reported  from  Minnesota. 

Atriplex  patula  L.  Sp.  PI.  1053.     1753. 

Coll.  :  Ballard  2577,  St.  Vincent;  2614,  2625,  Humboldt; 
2702,  Northcote ;  2760,  2771,  Hallock;  2722,  Kennedy; 
2772,  Warren;  2532,  Fergus  Falls. 
No  previous  authentic  collection  reported  from  Minnesota. 

Salicornia  herbacea  L.  Sp.  PI.  Ed.  2,  5.     1762. 

Coll.  :  Ballard  2680,  Northcote. 

"Growing  locally  along  the -drainage  from  a  salt  well.      No 
other  plants  found  growing  with  it."     Ballard. 

Dondia  depressa  (PURSH)  BRITTO.N  in  Britton  &  Brown,  111.  Fl. 

/.  /jpj.  i:  585.      1896. 

Coll.:  Ballard  2613,  2618,  Humboldt;  3701,  Northcote; 
2759,  Hallock;  2793,  Warren;  MacMillan  &  Skinner 
113,  Dugdale;  408,  Crookston. 

AMARANTHACE^E. 

Amaranthus  retroflexus  L.  Sp.  PL  991.     1753. 

Coll.  :  Ballard  2632,  St.  Vincent;  2707,  Northcote;  2717, 
Kennedy;  MacMillan  &  Skinner  321,  Crookston. 

Amaranthus  blitoides  S.   WATS.  Proc.   Am.   Acad.   12:  273. 

1877- 

Coll.  :  Ballard  2543,  Fergus  Falls;  MacMillan  &  Skinner, 
407,  Crookston. 

Amaranthus  graecizans  L.  Sp.  PL  990.     1753. 

Coll.  :  Ballard  2658,  St.  Vincent;  2603,  Humboldt;  2691, 
Northcote;  2718,  Kennedy. 


Wheeler:    THE  FLORA  OF  THE  RED  RIVER  VALLEY.         581 

NYCTAGINACE^E. 

Allionia  hirsuta  PURSH,  Fl.  Am.  Sept.  728.     1814. 
Coll.  :  MacMillan  &  Skinner  91,  Dugdale. 

CARYOPHYLLACE^E. 

Agrostemma  githago  L.  Sp.  PI.  435.     1753. 

Coll.  :   MacMillan  &  Skinner  125,  Dugdale. 

Silene  antirrhina  L.  Sp.  PI.  419.     1753. 

Coll.  :  MacMillan  &  Skinner  124,  Dugdale. 

Vaccaria  vaccaria  (L.)  BRITTON,  in  Britton  &  Brown  111.  Fl.  2  : 

18.     1897. 
Coll.  :   Ballard  2671,  St.  Vincent. 

Alsine  media  L.  Sp.  PI.  272.     1753. 
Coll.  :  Ballard  2736,  Kennedy. 

CERATOPHYLLACE^E. 

Ceratophyllum  demersum  L.  Sp.  PL  992.     1753. 
Coll.  :  Ballard  2654^,  St.  Vincent. 

RANUNCULACEjE. 

Actsea  alba  (L.)  MILL.  Card.  Diet.  Ed.  8,  No.  2.     1768. 
Coll.  :   MacMillan  &  Skinner  140,  Maple  lake. 

Anemone  cylindrica  A.  GRAY,  Ann.  Lye.  N.  Y.  3  :  221.     1836. 
Coll.  :   Ballard  2606,  Humboldt. 

Anemone  virginiana  L.  Sp.  PI.  540.     1753. 

Coll.  :  Ballard  2526,  Fergus  Falls  ;  MacMillan  &  Skinner 
27,  Crookston. 

Anemone  canadensis  L.  Syst.  Ed.  12,  3:  App.  231.     1768. 
Coll.  :  MacMillan  &  Skinner  39,  Crookston. 

Clematis  virginiana  L.  Amoen.  Acad.  4:  275.     1759. 
Coll.  :  MacMillan  &  Skinner  186,  Maple  lake. 

Ranunculus  scleratus  L.  Sp.  PI.  551.     1753. 

Coll.  :  Ballard  2644,  St.  Vincent;  2586,  Humboldt;  Mac- 
Millan &  Skinner  427,  Crookston. 

Ranunculus  pennsylvanicus  L.  f.  Suppl.  272.     1781. 

Coll.:  Ballard  2507,  Fergus  Falls;  2642,  St.  Vincent; 
2756,  Hallock  ;  MacMillan  &  Skinner  22,  Crookston ; 
288,  Maple  lake. 


582  MINNESOTA    BOTANICAL    STUDIES. 

Oxygraphis  cymbalaria  (PURSH)   PRANTL,  in  Engl.  &  Prantl, 

Nat.  Pfl.  Fam.  3:   Abt.  2,  63.      1891. 

Coll.:  Ballard    2508,    Fergus    Falls;   2645,   St.  Vincent; 
2788,  Warren;  MacMillan  &  Skinner  409,  Holmes. 

Thalictrum  purpurascens  L.  Sp.  PI.  546.     1753. 
Coll.  :   MacMillan  &  Skinner  32,  Crookston. 

BERBERIDACEjE. 

Caulophyllum  thalictroides  (L.)  MICHX.  Fl.  Bor.  Am.  i :  205. 

1803. 
Coll.  :  MacMillan  &  Skinner  286,  Thief  River  Falls. 

MENISPERMACE^:. 

Menispermum  canadense  L.  Sp.  PI.  340.     1753. 

Coll.:  Ballard  2661,   St.  Vincent;  MacMillan  &  Skinner 
194,  Crookston. 

CRUCIFER^E. 

Thlaspi  arvense  L.  Sp.  PI.  646.     1753. 

Coll.  :  Ballard  2573,  St.  Vincent;  2579,  Humboldt ;  2690, 
North  cote. 

Roripa  palustris  (L.)  BESS.  Enum.  27.     1821. 

Coll.  :  Ballard  2709,  Northcote ;  MacMillan  and  Skinner 
214,  Crookston. 

Roripa  hispida  (DESV.)  BRITTON,  Mem.  Torr.  Club,  5 :   169. 

1894. 
Coll.  :  Ballard  2639,  St.  Vincent. 

Sophia  hartwegiana  (FOURN.)  GREENE,  Pittonia,  3  :  95.     1896. 
Coll.  :  MacMillan  &  Skinner  363,  Crookston. 

Erysimum  cheiranthoides  L.  Sp.  PI.  661.     1753. 

Coll.  :  MacMillan  &  Skinner  5,  197,  Crookston. 

CRASSULACE.E. 

Penthorum  sedoides  L.  Sp.  PI.  432.     1753. 

Coll.:    MacMillan  &   Skinner   231,    Crookston;    Ballard 
2780,  Warren. 


Wheeler:    THE  FLORA  OF  THE  RED  RIVER  VALLEY.         583 

PARNASSIACE.E. 

Parnassia  caroliniana  Michx.  Fl.  Bor.  Am.  i :   184.     1803. 

Coll.  :  MacMillan  &  Skinner  94,  Maple  lake. 
Parnassia  palustris  L.  Sp.  PL  273.     1753. 

Coll.  :  Ballard  2512,  Fergus  Falls. 

ROSACES. 

Spiraea  salicifolia  L.  Sp.  PI.  489.     1753. 

Coll.  :    Ballard  2706,  Northcote ;    MacMillan   &  Skinner 
156,  Maple  lake. 

Rubus  strigosus  MICHX.  Fl.  Bor.  Am.   i :  297.     1803. 
Coll.  :  MacMillan  &  Skinner  3,  Crookston. 

Potentilla  leucocarpa  RYDBERG,  in  Britton  &  Brown,  111.  Fl.  2  : 

212.  f.  I Q2J..       1897. 

Coll.:  Ballard  2641,  St.  Vincent;  MacMillan  &  Skinner 
323,  Crookston. 

Potentilla  monspeliensis  L.  Sp.  PL  499.     1753. 

Coll.  :  Ballard  2515,  Fergus  Falls  ;  2728,  Kennedy  ;  Mac- 
Millan and  Skinner  72,  291,  Crookston. 

Potentilla  pennsylvanica  strigosa  PURSH,  FL  Am.  Sept.  356. 

1814. 

Coll.:    MacMillan   &   Skinner  40,    Crookston;    252   Red 
Lake  Falls  ;  374,  Holmes. 

Potentilla  effusa  DOUGL.  ;  Lehm.  Stirp.  Pug.  2:  8.     1830. 

Coll.  :  MacMillan  &  Skinner  385,  Holmes. 
No  previous  authentic  collection  reported  from  Minnesota. 

Argentina  anserina  (L.)  RYDBERG,  Mem.  Dept.  Bot.  Columbia 

Univ.  2  :   159.  pi.  98.      1898. 

Coll.  :   Ballard  2574,  St.  Vincent;  2626,  Humboldt ;  Mac- 
Millan &  Skinner  191,  221,  Crookston. 

Comarum  palustre  L.  Sp.  PL  502.     1753. 

Coll.  :  MacMillan  &  Skinner  143,  Maple  lake. 

Drymocallis  arguta  (PURSH)  RYDBERG,  Mem.  Dept.  Bot.  Co- 
lumbia Univ.  2  :   192.  pi.  102.      1898. 
Coll.  :  MacMillan  &  Skinner  90,  Dugdale  ;  251,  Crookston  ; 
377,  Holmes. 


584  MINNESOTA    BOTANICAL    STUDIES. 

Chamaerhodos  erecta  (L.)  BUNGE,  in  Ledeb.  Fl.  Alt.   i  :  430. 

1829. 

Coll.  :  MacMillan  &  Skinner  375,  Holmes. 
Not  previously  reported  from  Minnesota. 

Geum  virginianum  L.  Sp.  PI.  500.     1753. 

Coll.:  MacMillan  &  Skinner   213,  Crookston ;   250,  Red 
Lake  Falls;  369,  Holmes;  Ballard  2557,  St.  Vincent. 

Agrimonia   hirsuta    (MUHL.)    BICKNELL,    Bull.    Torr.    Club, 

23:509.     1896. 
Coll.  :   MacMillan  &  Skinner  354,  Shirley. 

Rosa  arkansana  PORTER,  Syn.  Fl.  Col.  38.     1874. 

Coll.:  Ballard   2681,  Northcote ;    MacMillan    &   Skinner 
328,  Crookston. 

POMACES. 

Crateegus  coccinea  L.  Sp.  PI.  476.     1753. 

Coll.  :   Ballard  2562,  St.  Vincent. 

The  determination  of  this  is  doubtful.      It  is  the  common  form 
in  northwestern  Minnesota  and  Manitoba. 

DRUPACE^:. 

Primus  americana  MARSH,  Arb.  Am.  in.     1785. 

Coll.  :   MacMillan  &  Skinner  192,  315,  Crookston. 

Prunus  serotina  EHRH.  Beitr.  3  :  20.     1788. 

Coll.  :  MacMillan  &  Skinner  168,  Maple  lake. 

PAPILIONACE^. 

Lotus  americanus  (NUTT.)  BISCH.  Litt.  Ber.  Linnasa,  14:   132. 

1840. 
Coll.:  Ballard  2771^,  Hallock. 

Psoralea  argophylla  PURSH,  Fl.  Am.  Sept.  475.     1814. 

Coll.:   Ballard   2608,    Humboldt;    MacMillan  &  Skinner 
49,  Crookston. 

Amorpha  fruticosa  L.  Sp.  PI.  713.     1753. 

Coll.  :  Ballard  2505,  Fergus  Falls  ;  MacMillan  &  Skinner 
160,  Maple  lake. 

Amorpha  nana  NUTT.  Fras.  Cat.     1813. 

Coll.  :  Ballard  2800,  Warren  ;   MacMillan  &  Skinner  356, 
Shirley. 


I 
Wheeler:   THE  FLORA  OF  THE  RED  RIVER  VALLEY-.        585 

Kuhnistera   Candida  (WILLD.)   KUNTZE,   Rev.   Gen.   PL    192. 

1891. 
Coll.  :  MacMillan  and  Skinner  247,  Red  Lake  Falls. 

Kuhnistera  purpurea  (VENT.)   MAcM.  Met.  Minn.  Val.  329. 

1892. 

Coll.:    Ballard   2610,   Humboldt ;    2765,   Hallock ;    Mac- 
Millan &  Skinner  62,  Crookston. 

Astragalus  carolinianus  L.  Sp.  PI.  757.     1753. 
Coll.  :  MacMillan  &  Skinner  66,  Crookston. 

Phaca  neglecta  T.  &  G.  Fl.  N.  A.  i :  344.     1838. 

Coll.  :  MacMillan  &  Skinner  149,  153,  Maple  lake. 

Glycyrrhiza  lepidota  PURSH,  Fl.  Am.  Sept.  480.     1814. 

Coll.  :  Ballard  2705,  Northcote  ;  MacMillan  &  Skinner  273, 
St.  Hilaire. 

Meibomia  canadensis  (L.)  KUNTZE,  Rev.  Gen.  PI.  195.     1891. 
Coll.  :  MacMillan  &  Skinner  85,  Maple  lake. 

Vicia  americana  MUHL.  ;  Willd.  Sp.  PL  3:  1096.     1803. 

Coll.  :   Ballard  2767,  Hallock;  MacMillan  &  Skinner  79, 
Dugdale. 

Falcata  comosa  (L.)  KUNTZE,  Rev.  Gen.  PL   182.     1891. 

Coll.:     MacMillan    &    Skinner   162,    Maple    lake;     212, 
Crookston. 

GERANIACEJE. 

Geranium  bicknellii  BRITTON,  Bull.  Torr.  Club,  24:  92.     1897. 
Coll.  :  MacMillan  &  Skinner  145,  Maple  lake. 

OXALIDACE^E. 

Oxalis  stricta  L.  Sp.  PL  435.     1753. 

Coll.  :  MacMillan  &  Skinner  428,  Crookston. 

LINAGES. 

Linum  sulcatum  RIDDELL,  Suppl.  Cat.  Ohio  PL  10.     1836. 
Coll.  :  MacMillan  &  Skinner  108,  Dugdale. 

RUTACE^E. 
Xanthoxylum   americanum   MILL.  Gard.  Diet.  Ed.  8,  no.    2. 

1768. 
Coll.  :  MacMillan  &  Skinner  157,  Maple  lake. 


586  MINNESOTA    BOTANICAL    STUDIES. 

EUPHORBIACEJE. 

Euphorbia  serpyllifolia  PERS.  Syn.  2  :   14.     1807. 

Coll.:   Ballard  2665,  St.  Vincent;   2682,  Northcote  ;   2774, 

Warren. 
Euphorbia  glyptosperma  ENGELM.  Bot.  Mex.  Bound.  Surv.  187 

1859. 
Coll.  :  Ballard  2604,  2605,  Hallock  ;   2734,  Kennedy. 

Euphorbia  maculata  L.  Sp.  PL  455.     1753. 
Coll.  :  Ballard  2664,  St.  Vincent. 

ANACARDIACE^. 

Rhus  glabra  L.  Sp.  PI.  265.     1753. 

Coll.  :  MacMillan  &  Skinner  195,  Crookston. 

ACERACE^:. 

Acer  saccharum  MARSH.  Arb.  Amer.  4.     1785. 

Coll.  :  MacMillan  &  Skinner  182,  Maple  lake. 

Acer  negundo  L.  Sp.  PL  1056.     1753. 

Coll.  :  MacMillan  and  Skinner  422,  Crookston. 

BALSAMINACE^:. 

Impatiens  biflora  WALT.  Fl.  Car.  219.     1788. 
Coll.  :  Ballard  2500,  Fergus  Falls. 

VITACE^E. 

Vitis  vulpina  L.  Sp.  PL  203.     1753. 

Coll.  :  MacMillan  &  Skinner  166,  Maple  lake. 


Tilia  americana  L.  Sp.  PL  514.     1753. 

Coll.  :  MacMillan  &  Skinner  320,  Crookston. 

HYPERICACE^:. 

Triadenum  virginicum  (L.)  RAF.  FL  Tell.  3  :  79.     1836. 
Coll.  :  MacMillan  &  Skinner  165,  Maple  lake. 

CISTACE^E. 

Lechea  stricta  LEGGETT  ;  Britton,  Bull.  Torr.  Club,  21  :  251. 

1894. 
Coll.  :  MacMillan  &  Skinner  no,  Maple  lake. 


Wheeler:   THE  FLORA  OF  THE  RED  RIVER  VALLEY.        587 

VIOLACE^E. 

Viola  obliqua  HILL,  Hort.  Kew.  316.  pi.  12.     1769. 

Coll.:  Ballard  2516,  Fergus    Falls;  MacMillan  &  Skin- 
ner 382,  Holmes. 

Viola  pedata  L.  Sp.  PI.  933.     1753. 

Coll.  :  MacMillan  &  Skinner  279,  Ives. 

Viola  canadensis  L.  Sp.  PI.  936.     1753. 

Coll.  :  MacMillan  &  Skinner  351,  Crookston. 


Elaeagnus  argentea  PURSH,  Fl.  Am.  Sept.   114.     1814. 

Coll.  :  Ballard  2578,  Humboldt  ;  2801,  Warren;  MacMil- 
lan &  Skinner  53,  Crookston. 

ONAGRACE^:. 

Epilobium  lineare  MUHL.  Cat.  39.     1813. 

Coll.  :  MacMillan  &  Skinner  292,  Crookston. 

Epilobium  coloratum   MUHL.;  Willd.   Enum.   i:  411.     1809. 
Coll.:    MacMillan  &  Skinner   115,    Dugdale  ;    116,  Maple 
lake;    244,    Red    Lake   Falls;    Ballard    2533,    Fergus 
Falls;   2631,  St.  Vincent;  2742,  Hallock. 

Onagra  biennis  (L.)  SCOP.  Fl.  Cam.  Ed.  2,  i  :  269.     1772. 
Coll.  :  MacMillan  &  Skinner  42,  Crookston. 

(Enothera  rhombipetala  NUTT.  ;  T.  &  G.  Fl.   N.  A.I:  493. 

1840. 
Coll.  :  Ballard  2749,  Hallock. 

Anogra  pallida  (LINDL.)  BRITTON,  Mem.  Torr.  Club,  5  :  234. 
^  1894. 
Coll.  :  MacMillan  &  Skinner  364,  Crookston. 

Meriolix  serrulata  (Nuxx.)  WALP.  Repert.  2  :  79.     1843. 

Coll.  :  MacMillan  &  Skinner  33,  Crookston;  109,  Mentor. 

Gaura  coccinea  PURSH,  Fl.  Am.  Sept.  733.     1814. 
Coll.  :  MacMillan  &  Skinner  430,  Holmes. 

HALORAGIDACE^:. 

Hippuris  vulgaris  L.  Sp.  PL  4.     1753. 

Coll.:   MacMillan  &  Skinner  230,  Crookston. 


588  MINNESOTA    BOTANICAL    STUDIES. 

Myriophyllum  spicatum  L.  Sp.  PL  992.     1753. 

Coll.  :  MacMillan  &  Skinner  393,  Holmes. 
Myriophyllum  verticillatum  L.  Sp.  PL  992.     1753. 

Coll.  :  Ballard  2653,  St.  Vincent. 

ARALIACEjE. 

Aralia  nudicaulis  L.  Sp.  PL  274.     1753. 

Coll.  :   MacMillan  &  Skinner  211,  Crookston. 

UMBELLIFER^E. 

Heracleum  lanatum  MICHX.  Fl.  Bor.  Am.  i :  166.     1803. 
Coll.  :   MacMillan  &  Skinner  202,  Crookston. 

Pastinaca  saliva  L.  Sp.  PL  262.     1753. 
Coll.  :  Ballard  2662,  St.  Vincent. 

Washingtonialongistylis(ToRR.)  BRITTON,  111.  Fl.  2  :  530.  1897. 
Coll.  :   MacMillan  &  Skinner  238,  Gentilly. 

Sium  cicutaefolium  GMEL.  Syst.  2  :  482.     1791. 

Coll.  :   Ballard  2593,  Humboldt. 
Zizia  aurea  (L.)  KOCH,  Nov.  Act.  Caes.  Leop.  Acad.  12  :   129. 

1824. 

Coll.  MacMillan  &  Skinner  164,   Maple  lake;   206,  327, 
Crookston. 

Zizia  cordata  (WALT.)  KOCH  in  DC.  Prodr.  4  :   100.     1830. 
Coll.  :   MacMillan  &  Skinner  253,  Red  Lake  Falls. 

Cicuta  bulbifera  L.  Sp.  PL  255.     1753. 

Coll.  :  MacMillan  &  Skinner  183,  Maple  lake. 

Deringa  canadensis  (L.)  KUNTZE,  Rev.  Gen.  PL  i :   266.    1891. 
Coll.  :   MacMillan  &  Skinner  208,.  325,  326,  Crookston. 

CORNACEvE. 

Cornus  stolonifera  MICHX.  Fl.  Bor.  Am.  i :  92.     1803. 
Coll.  :  MacMillan  &  Skinner  316,  Crookston. 

Cornus  candidissima  MARSH,  Arb.  Am.  35.     1785. 

Coll.  :  Ballard  2752,  Hallock ;  MacMillan  &  Skinner  242,, 
Red  Lake  Falls. 

ERICACEAE. 

Arctostaphylos  uva-ursi  (L.)  SPRENG.  Syst.  2  :  287.     1825. 
Coll.  :  MacMillan  &  Skinner  144,  Maple  lake. 


Wheeler :   THE  FLORA  OF  THE  RED  RIVER  VALLEY.        589 

VACCINIACE^E. 

Oxycoccus   oxycoccus  (L.)  MacM.   Bull.^  Torr.  Club,  19:15. 

1892. 
Coll.  :  MacMillan  &  Skinner  185,  Maple  lake. 

PRIMULACE^E. 

Steironema  ciliatum  (L.)  RAF.  Ann.  Gen.  Phys.  7 :  192.     1820. 
Coll.:  Ballard  2776,  Warren;  MacMillan  &  Skinner  16, 

Crookston. 
Steironema  lanceolata  (WALT.)  A.  GRAY,  Proc.  Am.  Acad.  12: 

63.     1876. 

Coll.  :  MacMillan  &  Skinner  98,  Maple  lake  ;  345,  Crook- 
ston. 

GENTIANACE^. 

Gentiana  detonsa  ROTTB.  Act.  Hafn.  10  :  254.     1770. 

Coll.  :  MacMillan  &  Skinner  184,  Maple  lake. 
Gentiana  acuta  MICHX.  Fl.  Bor.  Am.  i :  177.     1803. 

Coll.  :  MacMillan  &  Skinner  330,  346,  Crookston. 
No  previous  collections  from  Minnesota  in  the  Herbarium  of 

the  University. 
Gentiana  puberula  MICHX.  Fl.  Bor.  Am.  i :  176.     1803. 

Coll.  :  MacMillan  &  Skinner  54,  Crookston  ;  350  Shirley. 
Gentiana  andrewsii  GRISEB.   in  Hook.   Fl.   Bor.   Am.   2:   55. 

1834. 
Coll.  :  Ballard  2792,  Warren;  MacMillan  &  Skinner  181, 

Maple  lake. 

Gentiana  flavida  A.  GRAY,  Am.  Journ.  Sci.  (II)  i :  80.     1846. 
Coll. :  MacMillan  &  Skinner  81,  Dugdale. 

APOCYNACE.E. 

Apocynum  androsaemifolium  L.  S p.  PI.  213.     1753. 
Coll.  :  MacMillan  &  Skinner  179,  Maple  lake. 

ASCLEPIADACE^. 

Asclepias  incarnata  L.  Sp.  PI.  215.     1753. 

Coll.  :  MacMillan  &  Skinner  29,  Crookston ;  399,  Holmes. 

Asclepias  syriaca  L.  Sp.  PL  214.     1753. 

Coll.  :  MacMillan  &  Skinner  198,  Crookston. 


590  MINNESOTA    BOTANICAL    STUDIES. 

Asclepias  speciosa  TORR.  Ann.  Lye.  N.  Y.  2  :  218.     1826. 
Coll.  :  MacMillan'&  Skinner  58,  Crookston. 

CONVOLVULACE^E. 

Convolvulus  sepium  L.  Sp.  PI.  153.     1753. 

Coll.  :  Ballard  2566,  St.  Vincent ;  MacMillan  &  Skinner 
403,  Crookston. 

CUSCUTACE^E. 

Cuscuta  polygonorum  ENGELM.  Am.  Journ.  Sci.  43  :  342.    1842. 
Coll.  :  Ballard  2674,  St.  Vincent. 

Cuscuta  gronovii  WILLD.  ;  R.  &  S.  Syst.  6:   205.      1820. 

Coll.:  MacMillan   and     Skinner    223,     Crookston;     268, 
Thief  River  Falls  ;  376,  Holmes. 

BORAGINACEjE. 

Lappula  lappula  (L.)  KARST.  Deutsch.Fl.  979.     1880-83. 

Coll.  :  Ballard  2770,  Hallock  ;  MacMillan  &  Skinner  245, 
Red  Lake  Falls. 

Lappula  americana  (A.  GRAY)  RYDBERG,  Bull.  Torr.  Club,  24: 

294.     1897. 

Coll.  :  MacMillan  &  Skinner  243,  Red  Lake  Falls. 
No  previous  authentic  collection  reported  from    Minnesota. 
Previous  collections  of  this  species  in  this  State  have  been 
made  and  determined  as  L.  floribtmda  (Lehm.)  Greene. 

Onosmodium  carolinianum  (LAM.)  DC.  Prodr.  10  :  70.     1846. 
Coll.  :  MacMillan  &  Skinner  386,  Holmes. 

VERBENACE^. 

Verbena  hastata  L.  Sp.  PI.  20.     1753. 

Coll.:  Ballard  2502,  Fergus    Falls;    2647,   St.    Vincent; 
MacMillan  &  Skinner  30,  Crookston. 

LABIATJE. 

Teucrium  canadense  L.  S  p.  PI.  564.     1763. 

Coll.  :  MacMillan  &  Skinner  118,  Maple  lake. 

Scutellaria  lateriflora  L.  Sp.  PI.  598.     1753. 

Coll.  :  Ballard  2506,  Fergus  Falls ;   MacMillan  &  Skinner 
i,  Crookston. 


Wheeler:   THE  FLORA  OF  THE  RED  RIVER  VALLEY.        591 

Scutellaria  galericulata  L.  Sp.  PI.  509.     1753. 

Coll.:     MacMillan  &  Skinner  302,  Maple  lake. 
Agastache  anethiodora  (Nuxx.)  BRITXON,  in   Britton  &  Brown, 

111.  Fl.  3:85.     1898. 
Dracocephalum  parviflorum  NUTT.  Gen.  2  :  35.     1818. 

Coll.:  Ballard  2556,  St.  Vincent;  MacMillan  &  Skinner 

284  Thief  River  Falls. 
Physostegia  virginiana  (L.)   BENTH.    Lab.    Gen.   &  Sp.   504. 


Coll.:  Ballard   2564,81.   Vincent;   2766,  Hallock  ;  2773, 
Warren;  MacMillan  &  Skinner  21,  215,  Crookston. 

Galeopsis  tetrahit  L.  Sp.  PI.  579.     1753. 
Coll.  :  Ballard  2607,  Humboldt. 

Stachys  palustris  L.  Sp.  PI.  580.     1753. 

Coll.:    Ballard   2640,    2650,  St.    Vincent;    MacMillan   & 
Skinner  232,  Crookston. 

Monarda  fistulosa  L.  Sp.  PI.  22.     1753. 

Coll.  :  MacMillan  &  Skinner  248,  Red  Lake  Falls. 

Koellia  flexuosa  (WALT.)  MAcM.  Met.  Minn.  Val.  452.     1892. 
Coll.  :  MacMillan  &  Skinner  96,  Mentor. 

Lycopus  virginicus  L.  Sp.  PI.  21.     1753. 

Coll.  :  MacMillan  &  Skinner  161,  Maple  lake. 
Lycopus  americanus  MUHL.  ;  Bart.  Fl.  Phil.  Prodr.  15.     1815. 

Coll.  :  Ballard  2648,  St.  Vincent;   2737,  Hallock. 
Lycopus  lucidus  TURCZ.  ;  Benth.  in  DC.  Prodr.  12  :  178.     1848. 

Coll.  :  Ballard  2509,  Fergus  Falls. 
Mentha  canadensis  L.  Sp.  PI.  577.     1753. 

Coll.:  Ballard  2560,  St.  Vincent  ;   2592,  Humboldt  ;  Mac- 
Millan &  Skinner  4,  426,  Crookston. 

SOLANACE^:. 
Solanum  nigrum  L.  Sp.  PI.  186.     1753. 

Coll.  :  MacMillan  &  Skinner  201,  Crookston. 

SCROPHULARIACE^). 

Pentstemon  gracilis  NUTT.  Gen.  2:  52.     1818. 
Coll.  :   MacMillan  &  Skinner  80,  Maple  lake. 

Mimulus  ringens  L.  Sp.  PI.  634.     1753. 

Coll.  :    MacMillan  &  Skinner  224,  Crookston. 


592  MINNESOTA    BOTANICAL    STUDIES. 

Veronica  americana  SCHWEIN.  Benth.  in  DC.  Prodr.  10 :  468. 

1846. 
Coll.  :  Ballard  2744,  Hallock. 

Veronica  scutellata  L.  Sp.  PI.  12.     1753. 

Coll.  :  MacMillan  &  Skinner  395,  Holmes. 

Leptandra  virginica  (L.)  NUTT.  Gen.  i :  7.     1818. 

Coll.  :  MacMillan  &  Skinner  15,  Crookston. 
Gerardia  aspera  DOUGL.  ;  Benth.  in  DC.  Prodr.  10  :  517.    1846. 

Coll.  :  MacMillan  &  Skinner  36,  Crookston. 
Gerardia  tenuifolia  VAHL,  Symb.  Bot.  3:  79.     1794. 

Coll.:  Ballard  2510,  2536,  Fergus  Falls;   2748,  Hallock; 
MacMillan  &  Skinner  277,  Ives. 

Orthocarpus  luteus  NUTT.  Gen.  2:  57.     1818. 

Coll.:    Ballard  2688,  Northcote ;    2719,  Kennedy;    Mac- 
Millan &  Skinner  47,  Crookston. 

Pedicularis  lanceolata  MICHX.  Fl.  Bor.  Am.  2:  18.     1803. 

Coll.:     MacMillan    &    Skinner    93,    Maple    lake;     379, 
Holmes. 

PHRYMACE^E. 

Phryma  leptostachya  L.  Sp.  PI.  60 1.     1753. 

Coll.:    MacMillan    &    Skinner    240,  Gentilly ;    241,  Red 

Lake  Falls. 
Plantago  major  L.  Sp.  PI.  112.     1753. 

Coll.  :  Ballard  2535,  Fergus  Falls;   2667,  St.  Vincent. 
Plantago  eriopoda  TORR.  Ann.  Lye.  N.  Y.  2  :  237.     1827. 

Coll.  :   Ballard  2789,  Warren. 

RUBIACE.E. 

Houstonia  longifolia  GAERTN.  Fruct.  i :  226.     1788. 
Coll.  :  MacMillan  &  Skinner  274,  St.  Hilaire. 

Galium  boreale  L.  Sp.  PI.  108.     1753. 

Coll.  :  MacMillan  &  Skinner  289,  Maple  lake. 

Galium  trifidum  L.  Sp.  PL  105.     1753. 

Coll.  :  Ballard  2646,  St.  Vincent ;  MacMillan  &  Skinner 
314,  Crookston. 

CAPRIFOLIACE^E. 
Viburnum  opulus  L.  Sp.  PL  268.     1753. 

Coll.  :  MacMillan  &  Skinner  180,  Maple  lake. 


Wheeler:   THE  FLORA  OF  THE  RED  RIVER  VALLEY.         593 

Viburnum  lentago  L.  Sp.'Pl.  268.     1753. 

Coll.  :    MacMillan   &    Skinner    272,   Thief    River    Falls, 
317,  Crookston. 

Symphoricarpos  occidentalis  Hook.  Fl.  Bor.  Am.  i :  285.    1833. 
Coll.  :  Ballard  2750,  Hallock ;  MacMillan  &  Skinner  20, 
Crookston. 

Symphoricarpos  symphoricarpos  (L.)  MACM.  Bull.  Torr.  Club, 

19:   15.     1892. 
Coll.  :  Ballard  2550,  St.  Vincent. 

CUCURBITACE^E. 
Micrampelis  lobata  (MICHX.)  GREENE,  Pittonia,  2  :   128.     1890. 

CAMPANULACE.E. 

Campanula  rotundifolia  L.  Sp.  PL  163.     1753. 
Coll.  :  MacMillan  &  Skinner  236,  Gentilly. 

Campanula  aparinoides  PURSH,  Fl.  Am.  Sept.  159.     1814. 

Coll.:    MacMillan    &    Skinner    163,     Maple   lake;    383, 
Holmes. 

Lobelia  syphilitica  L.  Sp.  PI.  931.     1753. 

Coll.  :  Ballard  2503,  Fergus  Falls;  MacMillan  &  Skinner 
299,  Crookston. 

Lobelia  spicata  LAM.  Encycl.  3:  587.     1789. 

Coll.  :  MacMillan  &  Skinner  57,  297,  298,  Crookston. 

Lobelia  kalmii  L.  Sp.  PI.  930.     1753. 

Coll.  :  MacMillan  &  Skinner  296,  Maple  lake. 

CICHORIACE.E. 

Taraxacum  taraxacum  (L.)  KARST.  Deutsch.  Fl.   1138.     1880- 

83- 
Coll.:  Ballard  2669,  St.  Vincent;  MacMillan  &  Skinner 

413,  Crookston. 

Sonchus  arvensis  L.  Sp.  PI.  793.     1753. 
Coll.  :  Ballard  2596,  Humboldt. 

Sonchus  asper  (L.)  ALL.  Fl.  Ped.  i :  222.     1785. 
Coll.  :  MacMillan  &  Skinner  177,  Maple  lake. 

Lactuca  ludoviciana  (Nutt.)  DC.  Prodr.  7:  141.     1838. 
Coll.  :  MacMillan  &  Skinner  34,  Crookston. 


594  MINNESOTA    BOTANICAL    STUDIES. 

Lactucapulchella  (PURSH)  DC.  Prodr.  7:  134.     1838. 

Coll.  :  Ballard  2547,  St.  Vincent;  2617,  Humboldt ;  2712, 
Northcote  ;  2723,  Kennedy;  MacMillan  &  Skinner  31, 
Crookston. 

Agoseris  glauca  (PURSH)  GREENE,  Pittonia,  2  :  176.     1891. 

Coll.:  Ballard  2735,  Kennedy;  2791,  Warren;  Mac- 
Millan &  Skinner  43,  Crookston. 

Hieracium  canadense  MICHX.  Fl.  Bor.  Am.  2:  86.     1803. 

Coll.:  Ballard  2768,  Hallock  ;  2799,  Warren;  MacMillan 
&  Skinner  84,  Dugdale  ;  114,  Maple  Lake. 

Nabalus  albus  (L.)  HOOK.  Fl.  Bor.  Am.  i :  294.     1833. 

Coll.:  Ballard  2627,  St.  Vincent;  2781,  Warren;  Mac- 
Millan &  Skinner  76,  Dugdale. 

Nabalus  racemosus  (Micnx.)  DC.  Prodr.  7:  242.     1838. 

Coll.  :  Ballard  2559,  St.  Vincent;  2609,  Humboldt;  Mac- 
Millan &  Skinner  59,  Crookston;  263,  St.  Hilaire. 

AMBROSIACE^E. 

Iva  xanthifolia(FRESEN.)  NUTT.  Trans.  Am.  Phil.  Soc.  (II.)  7: 

347.     1841. 
Coll.  :  Ballard  2659,  St.  Vincent;   2716,  Kennedy. 

Ambrosia  trifida  L.  Sp.  PI.  987.     1753. 

Coll.  :  Ballard  2567,  St.  Vincent,  2704,  Northcote. 

Ambrosia  artemisiaefolia  L.  Sp.  PI.  987.     1753. 
Coll.  :  Ballard  2513,  Fergus  Falls. 

Ambrosia psilostachya  DC.  Prodr.  5:  526.     1836. 

Coll.  :  Ballard  '  2538,  Fergus  Falls  ;  2689,  Northcote  ; 
2790,  Warren;  MacMillan  &  Skinner  331,  Crookston. 

Xanthium  canadense  MILL.  Gard.  Diet.  Ed.  8,  no.  2.     1768. 
Coll.:  Ballard    2511,   Fergus    Falls;    2677,  St.  Vincent; 
2764,  Hallock;  MacMillan  &  Skinner  217,  Crookston. 

COMPOSITE. 

Vernonia  fasciculata  MICHX.  Fl.  Bor.  Am.  2  :  94.     1803. 
Coll.  :  MacMillan  &  Skinner  50,  Crookston. 

Eupatorium  maculatum  L.  Amoen.  Acad.  4:  288.     1755. 

Coll.:  MacMillan  &  Skinner  121,  Maple  lake;  391, 
Holmes. 


Wheeler:    THE  FLORA  OF  THE  RED  RIVER  VALLEY.         595 

Eupatorium  perfoliatum  L.  S p.  PI.  838.     1753. 

Coll.  :  Ballard  2522,  Fergus  Falls;  MacMillan  &  Skinner 
159,  Maple  lake. 

Laciniaria  punctata  (HooK.)  KUNTZE,  Rev.  Gen.  PI.  349.    1891 . 
Coll.  :  MacMillan  &  Skinner  92,  Dugdale. 

Laciniari  apycnostachya  (Micnx.)  KUNTZE,  Rev.  Gen.  PI.  349. 

1891. 
Coll.  :  MacMillan  &  Skinner  97,  Maple  lake. 

Laciniaria  scariosa  (L.)  HILL,  Veg.  Syst.  4  :  49.     1762. 

Coll.  :  Ballard  2797,  Warren;  MacMillan  &  Skinner  61, 
220,  Crookston. 

Grindelia  squarrosa  (PURSH)  DUNAL  in   DC.   Prodr.   5:    315. 

1836. 

Coll.:  Ballard  2554,  St.  Vincent;  2700,  Northcote  ;  2730, 
Kennedy  ;  MacMillan  &  Skinner  88,  Dugdale. 

Chrysopsis  hispida  (HooK.)  NUTT.  Trans.  Am.  Phil.  Soc.  (II.) 

7:  316.     1841. 

Coll.  :  MacMillan  &  Skinner  83,  Dugdale;  388,  Holmes. 
Not  previously  reported  from  Minnesota. 

Solidago  canadensis  L.  Sp.  PI.  878.     1753. 

Coll.:  Ballard  2517,  Fergus  Falls;  2565,  St.  Vincent; 
2711,  Northcote  ;  MacMillan  &  Skinner  303,  Crookston  ; 
397,  Holmes. 

Solidago  nemoralis  AIT.  Hort.  Kew.  3:  213.     1789. 
Coll.  :  MacMillan  &  Skinner  342,  344,  Crookston. 

Solidago  rigida  L.  Sp.  PI.  880.     1753. 

Coll.:  Ballard  2504,  Fergus  Falls;  2611,  Humboldt ; 
2683,  Northcote  ;  2731,  Kennedy  ;  2795,  Warren;  Mac- 
Millan &  Skinner  69,  339,  Crookston. 

Euthamia  graminifolia  (L.)  NUTT.  Gen.  2:  162.     1818. 
Coll.  :  Ballard  2521,  Fergus  Falls;  2796,  Warren. 

Aster  sagittifolius  WILLD.  Sp.  PI.  3:  2035.     1804. 

Coll.:  MacMillan  &  Skinner  25,  Crookston;  256,  St. 
Hilaire. 

Aster  novae-angliae  L.  Sp.  PI.  875.     1753. 

Coll.  :  Ballard  2779,  Warren  ;  MacMillan  &  Skinner  78, 
Dugdale. 


596  MINNESOTA    BOTANICAL    STUDIES. 

Aster  puniceus  L.  Sp.  PI.  875.     1753. 

Coll.  :   MacMillan  &  Skinner  287,  Thief  River  Falls. 

Aster  laevis  L.  Sp.  PI.  876.     1753. 

Coll.:  MacMillan  &  Skinner  190,  196,  311,  Crookston ; 
Ballard  2518,  Fergus  Falls;  2561,  2622,  St.  Vincent; 
2692,  Northcote ;  2738,  Hallock. 

Aster  sericeus  VENT.  Hort.  Cels.     1800. 

Coll.  :  MacMillan  &  Skinner  82,  Dugdale. 

Aster  ptarmicoides  (NEES)  T.  &  G.  Fl.  N.  A.  2 :   160.     1841. 
Coll.:   MacMillan  &  Skinner  60,  Crookston;   154,  Maple 
lake  ;  Ballard  2804,  Warren. 

Aster  salicifolius  LAM.  Encycl.  i :  306.     1783. 
Coll.  :  Ballard  2785,  Warren. 

Aster  paniculatus  LAM.  Encycl.  i :  306.     1783. 

Coll.:  Ballard  2568,  St.  Vincent ;  2674,  Northcote  ;  Mac- 
Millan &  Skinner  312,  Crookston. 

Aster  multiflorus  AIT.  Hort.  Kew.  3:  203.     1789. 

Coll.:  Ballard  2694,  Northcote;  2724,  Kennedy;  2802, 
Warren ;  MacMillan  &  Skinner,  336,  Crookston. 

Brachyactis  angustus  (LINDL.)  BRITTON,  in  Britton  &  Brown, 

111.  Fl.  3:  383.     1898. 
Coll.  :  Ballard  2545,  Fergus  Falls;   2784,  Warren. 

Erigeron  philadelphicus  L.  Sp.  PI.  863.     1753. 

Coll.:  MacMillan  &  Skinner  226,  360,  Crookston;  Bal- 
lard 2649,  St.  Vincent. 

Erigeron  ramosus  (WALT.)  B.S.P.  Prel.  Cat.  N.  Y.  27.     1888. 
Coll.:  MacMillan  &  Skinner  44,    Crookston;  95,   Maple 
lake. 

Leptilon  canadense  (L.)  BRITTON,  in  Britton  &  Brown  111.  Fl. 

3:  391.     1898. 

Coll.:  Ballard  2633,  St.  Vincent ;  MacMillan  &  Skinner 
37,  Crookston. 

Doellingeria  umbellata  pubens  (A.  GRAY)  BRITTON,  in  Britton  & 

Brown  111.  Fl.  3:  393.      1898. 
Coll.  :  MacMillan  &  Skinner  120,  Maple  lake. 

Heliopsis  scabra  DUNAL,  Mem.  Mus.  Paris,  5:  56.     1819. 

Coll.:  MacMillan  &  Skinner  117,  Maple  lake;  204, 
Crookston. 


Wheeler:   THE  FLORA  OF  THE  RED  RIVER  VALLEY.         597 

Rudbeckia  hirta  L.  Sp.  PL  907.     1753. 

Coll.:  Ballard  2563,  St.  Vincent ;  MacMillan  &  Skinner 
41,  Crookston. 

Rudbeckia  laciniata  L.  Sp.  PI.  906.     1753. 

Coll.:  Ballard  2660,  St.  Vincent ;  MacMillan  &  Skinner 
205,  Crookston. 

Ratibida  columnaris  (SiMs)  D.  DON;  Sweet,  Brit.  Fl.  Gard.  2: 

361.     1838. 

Coll.:  MacMillan  &  Skinner  75,  Maple  lake,  365, 
Holmes. 

Helianthus  annuus  L.  Sp.  PI.  904.     1753. 
Coll.  :  Ballard  2675,  St.  Vincent. 

Helianthus  scaberrimus  ELL.  Bot.  S.  C.  &  Ga.  2:  423.     1824. 
Coll.:   Ballard  2601,   Humboldt ;   2703,  Northcote ;   2739, 
Hallock ;  MacMillan  &   Skinner  333,  Crookston;  367, 
Holmes. 

Helianthus  maximiliani  SCHRAD.  Ind.  Sem.  Hort.  Goett.    1835. 
Coll.:  Ballard  2572,  St.  Vincent ;   2696,  Northcote  ;   2727, 
Kennedy;  MacMillan  &  Skinner  352,  353,  Shirley. 

Helianthus  grosse-serratus  MARTENS,  Sel.  Sem.  Hort.  Loven. 

^  1839. 

Coll.:  MacMillan  &  Skinner  295,  429,  Crookston;  378, 
Holmes. 

Helianthus  tuberosus  L.  Sp.  PI.  905.     1753. 

Coll.  :  MacMillan  &  Skinner  203,  Crookston  ;  392,  Holmes. 

Bidens  Isevis  (L.)  B.S.P.  Prel.  Cat.  N.  Y.  29.     1888. 

Coll.:  MacMillan  &  Skinner  264,  St.  Hilaire ;  362, 
Crookston;  371,  Holmes. 

Bidens  cernua  L.  Sp.  PI.  832.     1753. 

Coll.  :  MacMillan  &  Skinner  9,  Crookston. 

Bidens  frondosa  L.  Sp.  PI.  832.     1753. 

Coll.:  Ballard  2623,  Humboldt;  2678,  St.  Vincent ;  2710, 
Northcote;  2733,  Kennedy;  2763,  Hallock;  MacMil- 
lan &  Skinner  n,  38,  207,  Crookston. 

Helenium  autumnale  pubescens  (AiT.)  BRITTON,   Mem.  Torr. 

Club,  5:  339.     1894. 
Coll.  :  MacMillan  &  Skinner  146,  Maple  lake. 


598  MINNESOTA    BOTANICAL    STUDIES. 

Gaillardia  aristata  PURSH,  Fl.  Am.  Sept.  573.     1814. 

Coll.  :   MacMillan  &  Skinner  380,  Holmes. 
No  previous  authentic  collections  from  Minnesota  in  the  Her- 
barium of  the  University. 

Achillas  millefolium  L.  Sp.  PI.  899.     1753. 

Coll.:  Ballard  2624,  St.  Vincent;  MacMillan  &  Skinner 
45,  Crookston. 

Artemisia  caudata  MICHX.  Fl.  Bor.  Am.  2:  129.     1803. 
Coll.  :  MacMillan  &  Skinner  73,  Crookston. 

Artemisia  dracunculoides  PURSH,  Fl.  Am.  Sept.  742.     1814. 
Coll.  :   MacMillan  &  Skinner  65,  Crookston. 

Artemisia  frigida  WILLD.  Sp.  PL  3:  1838.     1804. 

Coll.  :  MacMillan  &  Skinner  74,  Crookston ;  Ballard 
2597,  Humboldt. 

Artemisia  absinthium  L.  Sp.  PL  848.     1753. 
Coll.  :  Ballard  2553,  St.  Vincent. 

Artemisia  biennis  WILLD.  Phytogr.  n.     1794. 
Coll.  :  Ballard  2726,  Kennedy. 

Artemisia  gnaphalodes  NUTT.  Gen.  2  :  143.     1818. 

Coll.:  Ballard  2619,  Humboldt;  2698,  Northcote ;  2732, 
Kennedy;  2798,  Warren;  MacMillan  &  Skinner  55, 
Crookston. 

Arctium  lappa  L.  Sp.  PL  816.     1753. 

Coll.  :  MacMillan  &  Skinner  210,  Crookston. 

Carduus  altissimus  L.  Sp.  PL  824.     1753. 

Coll.  :  Ballard  2501,  Fergus  Falls;  MacMillan  &  Skinner 
170,  Maple  lake. 

Carduus  discolor  (MUHL.)  NUTT.  Gen.  2:  130.     1818. 
Coll.  :  MacMillan  &  Skinner  12,  Crookston. 

Carduus  undulatus  NUTT.  Gen.  2  :  130.     1818. 
Coll.  :  MacMillan  &  Skinner  48,  Crookston. 

Carduus  arvensis  (L.)  ROBS.  Brit.  FL  163.     1777. 

Coll.:  Ballard  2548,  St.  Vincent ;  MacMillan  &  Skinner 
283,  Thief  River  Falls. 


Wheeler:   THE  FLORA  OF  THE  RED  RIVER  VALLEY.        699 

EXPLANATION  OF  PLATK  XXXIV. 

General  view  of  prairie  near  Shirley,  Minn.  This  is  the  charac- 
teristic aspect  of  mesophytic  prairie  in  the  Red  River  valley.  The 
shrubs  are  Salix  humilis,  for  the  most  part.  The  herbs  in  the  fore- 
ground are  Asters.  The  view  shows  a  minor  tension  line  in  which 
Junciis  dudleyi\%  an  abundant  plant.  On  the  right  a  growth  of  Poly- 
gonum  intermixed  with  Andropogon  is  seen.  The  view  gives  an  idea 
of  the  variety  of  the  prairie  vegetation. 

PLATE  XXXV. 

An  island  of  Hordcum  surrounded  by  a  border  zone  of  Salix  inter- 
mingled with  Symphoricarpos  and  Solidago.  Such  circular  patches  of 
squirrel-tail  grass  marking  slight  depressions  in  the  prairie  are  not  un- 
common and  often  reach  a  considerable  size, even  covering  several  acres. 

PLATE  XXXVI. 

Prairie  near  Gentilly,  Minn.  In  the  background  is  seen  the  shrubby 
and  scanty  arboreal  vegetation  along  the  Red  Lake  river.  In  the 
middle  distance  a  minor  tension  line  of  Hordeum  is  apparent  extending, 
in  this  case,  several  miles  along  the  river.  In  the  foreground  Nabalus 
racemosus,  a  characteristic  wand  plant  of  the  region,  is  seen  forming 
an  almost  circular  patch  in  the  general  grass  vegetation. 

PLATE  XXXVII. 

Gopher  mound  with  characteristic  vegetation.  These  mounds  made 
by  Geomys  bursarius  are  abundant  on  the  prairie  throughout  the  dis- 
trict. Somewhat  more  xerophytic  plants  inhabit  them  than  are  found 
upon  the  level  prairie  where  they  occur.  Upon  this  particular  mound 
hazel-brush,  Artemisia,  Bouteloua,  Solidago  rigida  and  other  semi- 
xerophytic  or  strongly  xerophytic  plants  have  secured  a  foothold. 

PLATE  XXXVIII. 

A  growth  of  silver-berry — Eleagnus  argentea.  This  plant  is 
abundant  throughout  the  district  studied,  in  dry  declivities  or  on  slopes 
of  the  rolling  prairie.  It  is  also  abundant  in  pastures  along  the  Red 
Lake  river. 

PLATE  XXXIX. 

View  of  the  margin  of  a  grass  meadow  in  the  poplar  scrub  near 
Maple  lake  showing  three  stands  of  Salix  lucida,  a  common  plant  of 
the  tension  zone  between  the  meadow  and  the  scrub. 

PLATE  XL. 

View  of  xerophilous  vegetation  on  knolls  along  a  coulee  cut  in  the 
raised  beach  of  the  extinct  Lake  Agassiz  near  Fertile,  Minn.  The 


600  MINNESOTA    BOTANICAL    STUDIES. 

brows  of  the  knolls  are  occupied  almost  exclusively  by  an  Artemisia 
formation  in  which  three  or  four  species  are  present,  Artemisia 
frigida  being  the  most  abundant.  Scrub  poplars,  hazel  and  Quercus 
form  a  sparse  "gallery  wood."  On  the  upland  Gaillardia,  Amor- 
pha,  Gaura  and  other  xerophytic  herbs  and  shrubs  of  the  prairie  are 
abundant. 

PLATE  XLI. 

Elm  woods  along  the  Red  Lake  river  near  Crookston,  Minn.  The 
bottoms  being  subject  to  overflow,  show  a  scanty  undergrowth  mostly 
herbaceous,  though  with  a  few  shrubs  of  Ribes,  Rubus  and  Corylus. 
In  such  glades  the  bolls  of  the  trees  are  commonly  distorted  and 
scarred  owing  to  the  battering  which  they  receive  when  young  by 
driftwood  and  flotsam  during  times  of  high  water. 


VOL.  II.  MINNESOTA  BOTANICAL  STUDIES.  PART  V. 


PLATE  XXX IV. 


VOL.11.  MINNESOTA  BOTANICAL  STUDIES.  PARTY. 


PLATE  XXXV. 


)L.  II. 


MINNESOTA  BOTANICAL  STUDIES. 


PLATE  XXXVI. 


VOL.  II. 


MINNESOTA  BOTANICAL  STUDIES. 


PART  V. 


PLATE  XXXVII. 


VOL.  II. 


MINNESOTA  BOTANICAL  STUDIES. 


PART  V. 


PLATE  XXXVIII. 


VOL.11.  MINNESOTA  BOTANICAL  STUDIES.  PARTY. 


PLATE  XXXIX. 


VOL.11.  MINNESOTA  BOTANICAL  STUDIES.  PARTY. 


PLATE  XL 


VOL.  II. 


)TA  BOTANICAL  STUDIES- 


PART  V. 


PLATE  XLI. 


XXXIII.  OBSERVATIONS   ON    Gigartina  exasperata 

HARV. 


H.   B.   HUMPHREY. 


The  plants  used  in  the  preparation  of  this  paper  were  col- 
lected by  Miss  Josephine  E.  Tilden,  in  Puget  Sound  near  Seattle, 
Washington,  in  August,  1897. 

They  are  found  growing  at  a  depth  of  six  fathoms  though  thriv- 
ing in  shallower  water.  In  July,  1898,  several  plants  were  col- 
lected near  Tracyton,  Washington,  at  a  depth  of  about  four 
fathoms  attached  to  rocks  in  quiet  waters.  These  plants  were 
generally  large  and  well  developed  and  were  somewhat  loosely 
attached  to  \\$  substratum.  Their  position  in  the  water  was 
erect  except  in  certain  places  where  a  tidal  current  was  present. 
Plants  found  in  localities  washed  by  swift  tidal  currents  were 
smaller,  thicker  and  more  firmly  attached  to  the  substratum. 

The  material  was  preserved  in  alcohol,  consequently  the  plant 
could  not  be  studied  in  its  natural  condition.  All  sections  were 
cut  by  means  of  a  freezing  microtome.  Material  imbedded  in 
gelatin  when  sectioned  proved  useless  as  the  cells  were  swollen 
to  such  a  degree  as  to  appear  unnatural.  Portions  of  the  frond 
were  then  sectioned  directly  from  the  alcoholic  solution  with 
good  results. 

The  stains  employed  were  Delafield's  hasmatoxylin,  methyl 
blue,  methyl  violet,  iodine  and  fuchsin.  Delafield's  hsematoxy- 
lin  proved  a  good  nuclear  stain.  Methyl  blue  was  used  in  stain- 
ing cell  walls  but  was  not  as  satisfactory  as  methyl  violet. 
Iodine  was  used  in  staining  carpospores  and  brings  out  very 
clearly  the  distinction  between  them  and  surrounding  tissue. 
Fuchsin  proved  a  very  satisfactory  stain  used  in  connection  with 
the  study  of  protoplasmic  pits,  coloring  them  a  deep  red. 

Sections  were  all  treated  with  staining  solution  and  then 
mounted  directly  in  glycerine  jelly,  making  a  permanent  mount. 

601 


602  MINNESOTA    BOTANICAL    STUDIES. 

Holdfast:  The  holdfast  is  a  disc-shaped  organ,  exhibiting 
considerable  variation  in  size.  In  the  case  of  a  single  frond  the 
holdfast  is  not  much  greater  in  diameter  than  the  stipe  immedi- 
ately above.  The  under  surface  is  smooth  and  somewhat  flex- 
ible, though  in  comparison  with  other  tissues  it  shows  greater 
rigidity  and  strength.  It  is  not  unusual  to  find  several  fronds 
attached  to  one  common  holdfast  which,  upon  close  examina- 
tion, presents  the  appearance  of  a  compound  organ,  in  some 
instances  measuring  nine  mm.  in  diameter. 

The  tissue  of  the  holdfast  is  unlike  that  of  any  other  part  of 
the  plant.  PI.  42,  Fig.  2,  represents  a  longitudinal  section 
through  a  portion  of  the  holdfast  showing  distinctive  areas  from 
the  point  of  attachment  to  the  substratum  to  the  tissue  of  the 
stipe.  It  was  found  that  in  removing  the  plant  from  its  point 
of  attachment  the  cuticle  was  removed  from  the  holdfast  leaving 
exposed  those  cells  immediately  adjacent,  represented  by  («). 
These  cells  appear  to  be  somewhat  irregular  in  outline,  though 
generally  quadrilateral,  and  are  characterized  by  their  exceed- 
ingly thick  walls.  Approaching  the  stipe  these  cells  are  slightly 
modified  and  in  conjunction  with  them  are  found  rather  long 
somewhat  egg-shaped  cells,  densely  filled  with  coritents.  These 
cells,  along  with  the  others,  are  arranged  approximately  in  rows 
extending  vertically  through  the  holdfast.  Protoplasmic  connec- 
tion exists  between  all  cells  and  the  cell  arrangement  is  so  com- 
pact as  to  give  great  strength  and  rigidity  to  the  tissue.  These 
cells  (PI.  42,  Fig.  2,  d)  are  slightly  modified  and  in  conjunc- 
tion with  small,  somewhat  spherical  cells  closely  attached  and 
densely  filled  with  granular  contents.  Abutting  upon  this  area 
are  the  filamentous  cells  of  the  stipe,  which  are  very  similar  to 
those  found  elsewhere  in  the  frond. 

Stipe. — In  the  early  stages  of  the  plant's  growth  the  stipe 
is  hardly  to  be  distinguished  from  the  lamina,  but  as  the  frond 
reaches  maturity  the  stipe  becomes  a  well-marked  organ  of 
deep  red  color.  Immediately  above  the  holdfast  it  is  circular, 
but  as  it  gradually  merges  into  the  lamina  it  loses  its  charac- 
teristic shape,  becoming  much  expanded  in  one  diameter  and 
thinner  in  the  other.  The  stipe  seldom  exceeds  a  length  of  20 
mm.,  while  the  diameter  varies  from  2  to  5  mm. 

The  stipe  exhibits  a  structure  similar  to  that  of  the  lamina, 
though  in  the  former  the  cells  possess  shorter  diameters  and 
the  arrangement  is  more  compact,  thus  affording  greater  rigid- 


Humphrey :  OBSERVATIONS  ON  Gigartina  exasperata  HARV.     603 

ity  and  strength.  The  epidermal  cells  throughout  the  entire 
plant  are  enveloped  by  a  firm  cellulose  sheath  of  variable 
thickness,  from  three  to  ten  mic.  (PL  42,  Fig.  4,  a.)  This 
cuticle  is  somewhat  elastic,  smooth  and  highly  transparent. 
PI.  42,  Fig.  5,  a  and  6,  represent  surface  views  of  a  portion 
of  the  frond,  showing  epidermal  cells  as  seen  through  the 
overlying  cuticle.  By  focusing,  deeper  cells  beneath  the  epi- 
dermal layer  may  be  seen. 

Beneath  the  epidermal  cells  and  in  connection  with  them  are 
the  pseudo-cortical  cells,  presenting  an  almost  spherical  outline 
and  a  somewhat  loose  though  definite  arrangement.  These 
cells  as  well  as  the  epidermal  ones  are  densely  filled  with  pro- 
toplasmic contents,  though  unlike  the  epidermal  cells  they  con- 
tain no  chromatophores. 

The  sections  of  the  stipe  were  stained  with  an  alcoholic  so- 
lution of  methyl  blue  which  gave  a  very  satisfactory  cellulose 
reaction  and  revealed  the  fact  that  all  the  cells  were  imbedded 
in  a  dense  gelatinous  matrix  between  which  and  the  cell  walls  it 
is  not  easy  to  distinguish. 

Adjoining  the  pseudo-cortical  cells  and  occupying  the  cen- 
tral region  of  the  stipe  is  the  pseudo-medullary  area  composed 
of  irregular  cells.  PL  42,  Fig.  3,  and  PL  42,  Fig.  4,  represent 
transverse  and  longitudinal  sections  of  the  stipe.  In  PL  42, 
Fig.  3,  c,  a  network  of  somewhat  filamentous  cells  is  seen  to 
be  interwoven  with  other  cells  of  different  form  forming  alto- 
gether a  rather  loose  arrangement. 

Lamina. — The  general  shape  of  the  lamina  is  almost  in- 
variably cuneate,  attaining  its  greatest  diameter  a  little  way 
from  the  apex.  In  all  cases  the  frond  is  flat  and  not  greater  than 
three  mm.  in  thickness,  and  when  dry  is  quite  translucent. 

It  commonly  grows  from  30  to  50  cm.  in  length  and  from  6 
to  18  cm.  in  width,  thus  showing  considerable  variation  in  size. 
In  shape  it  is  quite  as  variable  ;  some  fronds  being  branched  pro- 
fusely while  others  show  little  or  no  branching  whatever. 

New  fronds  arise  from  the  base  of  the  stipe  forming  at  first 
somewhat  club-shaped  or  pointed  bodies,  but  later  expand  and 
assume  the  characteristic  shape  of  the  mature  frond.  (PL  42, 
Fig.  i.)  Both  sides  of  the  frond,  including  the  margin,  are 
thickly  studded  with  cystocarps  and  numerous  epidermal  prolif- 
erations. Near  the  base  of  the  frond  on  each  side  is  a  small 
area  totally  void  of  proliferations.  Here  the  frond  is  thicker 


604  MINNESOTA    BOTANICAL    STUDIES. 

than  elsewhere,  more  deeply  colored  and  possesses  a  glossy 
smoothness. 

The  cystocarps  sometimes  appear  as  surface  elevations 
though  commonly  they  are  developed  in  the  marginal  and  sur- 
face proliferations.  They  are  most  numerous  and  attain 
greatest  size  in  the  marginal  area  while  at  the  center  they  are 
scattering  and  poorly  developed,  numbering  from  8  to  10  per 
sq.  cm.  as  compared  with  15  to  18  near  the  margin. 

^The  broad  flat  branches  of  the  lamina,  owing  to  extreme 
thinness  and  position,  bear  few  cystocarps  though  the  number  of 
proliferations  may  be  great. 

The  epidermal  cells  of  the  lamina  are  very  similar  in  every 
respect  to  those  of  the  stipe  except  that  the  arrangement  is  less 
compact.  The  same  may  be  said  regarding  the  pseudo-cortical 
area,  but  a  difference  is  seen  in  the  pseudo-medullary  cells  ; 
these  are  all  filamentous,  densely  rilled  with  granular  proto- 
plasmic contents  (PI.  42,  Figs.  6  and  7)  and  so  joined  as  to  form 
a  complete  network. 

Through  the  use  of  certain  staining  reagents  it  was  found  that 
a  protoplasmic  connection  existed  between  the  several  cells  of 
the  frond,  best  seen  in  the  pseudo-medullary  region  of  the 
lamina.  (PI.  42,  Fig.  8.)  On  further  examination,  using  alco- 
holic solution  of  fuchsin  as  a  staining  reagent,  protoplasmic 
pits  were  seen  to  exist  between  the  several  cells.  These  pits 
were  composed  in  every  case  of  two  minute  callous  plates  which 
when  stained  were  found  to  give  a  reaction  similar  to  that  of  pro- 
toplasm. It  was  not  possible  to  determine  the  function  of  these 
connections,  but  no  doubt  they  serve  as  paths  of  communication 
between  cells.  In  Schmitz's  discussion  of  the  protoplasmic 
pits  he  shows  that  they  are  traversed  by  plasma-cords  which 
serve  for  conduction  of  dynamic  influences  from  cell  to  cell. 
He  believes  a  transfer  of  dissolved  food  material  possible  be- 
cause of  the  pores  in  the  pit,  but  does  not  regard  as  probable 
the  transfer  of  protoplasm. 

Proliferations  and  cystocarp. — PI.  42,  Fig.  9,  represents  an 
early  stage  in  the  development  of  a  proliferation.  Certain  of 
the  epidermal  cells  become  slightly  modified  in  shape,  cell  di- 
vision takes  place  vertically  and  apparently  transversely.  This 
increase  in  the  number  of  cells  causes  an  elevation  to  develop 
and  as  it  continues  a  well-developed  proliferation  eventually  pre- 
vails which  may  or  may  not  bear  a  cystocarp. 


Humphrey :  OBSERVATIONS  ON  Gigartina  exasperata  HARV.     605 

In  the  material  at  hand  the  writer  was  unable  to  secure  any 
sections  showing  tetraspores. 

The  development  of  the  cystocarp,  however,  was  quite  clearly 
brought  out.  As  the  proliferation  advances  in  its  development 
there  arises  an  irregular  cellular  formation  of  gonimoblast  fila- 
ments and  sterile  tissue  in  the  interior  of  which  groups  of 
branched  filaments  develop  the  carpospores.  This  entire  for- 
mation is  surrounded  by  a  definite  area  of  cells  forming  the 
cystocarpic  wall.  As  the  cystocarp  advances  towards  maturity 
a  perforation  occurs  through  the  breaking  down  and  gradual 
dissolution  of  certain  cells,  thus  furnishing  the  mature  spores  an 
avenue  of  escape  (PL  42,  Fig.  u).  The  tissue  of  the  pro- 
liferation surrounding  the  spore  cavity  is  similar  to  that  of  the 
lamina  proper,  except  that  the  cells  are  more  compactly  ar- 
ranged. 

Sections  of  this  tissue  were  treated  with  Delafield's  haema- 
toxylin  which  proved  to  be  a  good  nuclear  stain  revealing  in 
several  cells  well-marked  nuclei.  (PI.  42,  Fig.  12.) 

Several  sections  were  made  in  order  to  determine  the  struc- 
ture and  characteristics  of  the  mature  cystocarp.  PL  42, 
Fig.  13,  represents  such  a  cystocarp  showing  the  spores  ar- 
ranged in  groups  surrounded  by  apparently  empty  filamentous 
cells,  thus  forming  a  compound  cystocarp.  Previous  to  the 
maturity  of  the  spores  they  are  all  attached  to  the  gonimoblastic 
filaments  of  which  they  were  originally  a  part.  They  are  evi- 
dently attached  by  means  of  protoplasmic  threads,  though  no 
evidence  of  pits  occurred.  (PL  42,  Fig.  15.)  The  carpo- 
spores, when  mature,  measure  from  10  to  12  /*  along  one  di- 
ameter and  u  to  13  fj.  along  the  other,  while  the  cystocarp 
measures  from  two  to  three  mm.  in  diameter  through  the  con- 
ceptacle. 

BIBLIOGRAPHY. 

Harvey,  W.  H.  Notice  of  a  collection  of  algae  made  on  the  north- 
west coast  of  North  America,  chiefly  at  Vancouver's  Island,  by 
David  Lyall  in  1859-61.  (Jour.  Proc.  of  Linn.  Soc.  6:  172, 
173.  1862. 

Cocks,  J.  Observations  on  the  growth  and  time  of  appearance  of 
some  of  the  marine  algae.  Jour.  Proc.  of  Linn.  Soc.  4:  101- 
106.  1860. 


606  MINNESOTA    BOTANICAL    STUDIES. 

Stromfelt,  H.  F.  G.  Untersuchungen  iiber  die  Haftorgane  der  Algen. 
Bot.  Cent.  33:  381,  382,  395-400.  iSSS.  Hedwigia,  27:  143. 
1888. 

Harvey,  W.  H.  Characters  of  new  algae  chiefly  from  Japan  and  ad- 
jacent regions.  Proc.  Am.  Acad.  4:  332.  1857-1860. 

Schmitz,  Fr.  Untersuchungen  iiber  die  Befruchtung  der  Florideen. 
Bot.  Zeit.  41  :  608-613,  145.  1883. 

Schmitz,  Fr.  Knollchenartige  Auswuchse  an  den  sprozzen  einiger 
Florideen.  Bot.  Zeit.  38:  624.  1892. 

Mdbius,  M.  Morphologic  der  haarartigen  Organe  bei  den  Algen. 
Biol.  Cent.  12:  73.  1892. 

Jonsson,  B.  Beitrage  zur  Kenntniss  des  Dickenzuwachses  der  Rho- 
dophyceen.  Lunds.  Univers.  Aarskr.  27:  41.  2  pi.  1890-91. 

Wille,  N.  Beitrag  zur  Entwicklungsgeschichte  der  physiologischen 
Gewebsysteme  bei  einigen  Florideen.  Nova  Acta.  Acad.  Leop. 
Carol.  52  :  49-100.  pL  3-8.  1887. 

Osterhout,  W.  J.  V.  A  Simple  Freezing  Device.  Bot.  Gaz.  21  : 
195-201.  1896. 

Moore,  L.  Le  M.  Studies  in  Vegetable  Biology — Observations  on  the 
Continuity  of  Protoplasm.  Journ.  Linn.  Soc.  21  :  595-621.  pi. 
19-21.  1886. 

DESCRIPTION  OF  PLATE  XLII. 

Figure  i.  A  typical  frond  of  Gigartina  exasperata  one-fourth 
natural  size. 

Figure  2.   Longitudinal  section  of  holdfast.      x  450. 

Figure  3.  Cross  section  of  stipe,  a,  Epidermal  cells;  b,  pseudo- 
cortical  cells;  c,  pseudo-medullary  cells.  x  450. 

Figure  4.  Longitudinal  section  of  stipe,  a,  Epidermal  cellulose 
sheath.  x  450. 

Figure  5.  (a)  Surface  view  of  frond,  showing  epidermal  cells 
through  the  transparent  epidermal  sheath.  (<$)  Surface  view  of  frond 
showing  cells  beneath  the  epidermal  cells.  X  900. 

Figure  6.  Cross  section  of  frond.      x  450. 

Figure  7.  Filamentous  cells  of  the  pseudo-medullary  area  showing 
granular  contents.  x  900. 


Humphrey :  OBSERVATIONS  ON  Gigartina  exasperata  HARV.     607 

Figure  8.  Protoplasmic  connections  between  cells  of  pseudo-medul- 
lary area.      x  900. 

Figure  9.  An  early  stage  in  development  of  a  proliferation  showing 
growing  point  (a).       X  450. 

Figure  10.  A  portion  of  the  margin  of  a  frond  showing  prolifera- 
tions and  cystocarps. 

Figure  1 1.   Longitudinal  section  of   mature  cystocarp.       O,  carpo- 
stome;  a,  conceptacle. 

Figure  12.  Portion  of  tissue  surrounding  a  conceptacle  showing  cell 
nuclei.      X  7°°' 

Figure  13.   Cross  section  of  mature  cystocarp  showing  spores. 

Figure  14.  Two  groups  of  spores  separated  by  elongated  sterile 
cells.      X  150. 

Figure  15.    Immature  carpospores   still   attached    to  the  gonimo- 
blastic  filaments;   («)  a  spore  separating  from  filaments.       X48o. 


VOL.  II. 


MINNESOTA  E 


i 


-    .  ,. 


3 


ANICAL  STUDIES. 


PART  V 


XLII 


NG   CO.,    BOSTON. 


XXXIV.  OBSERVATIONS   ON   THE   ALG^   OF  THE 
ST.    PAUL   CITY   WATER. 


M.  G.  FANNING. 


St.  Paul  receives  its  water  supply  from  twenty-two  lakes 
north  of  the  city  which  are  situated  on  both  sides  of  a  divide  or 
watershed.  The  area  from  which  the  water  is  received  extends 
about  twenty  miles  north  of  the  city.  The  greater  part  of  the 
water  comes  immediately  from  Lake  Vadnais,  which  in  return 
receives  its  supply  from  chains  of  lakes  through  brooks,  artifi- 
cial canals,  conduits,  etc.  As  these  lakes  are  separated  by  a 
divide  pumping  stations  are  provided  at  Centreville  lake  and 
Baldwin's  lake  to  force  the  water  over  the  divide.  Besides  the 
lakes,  groups  of  artesian  wells  add  to  the  supply  and  help  to 
lower  the  temperature  of  the  water  during  the  summer  months. 
There  are  nine  wells  at  Lake  Vadnais  and  twenty-eight  at  Cen- 
treville lake,  making  thirty-seven  in  all  with  depths  varying 
from  sixty-three  to  eight  hundred  and  sixty-five  feet. 

Pleasant  lake  receives  the  water  from  the  north  slope  of  the 
watershed  ;  from  here  the  water  flows  from  Lake  Vadnais,  then 
it  is  conveyed  four  and  one-half  miles  through  a  conduit  to  the 
pumping  station.  The  elevated  portions  of  the  city  receive  the 
water  directly  from  the  pumping  station.  Other  parts  are  sup- 
plied by  gravity  with  water  from  Lakes  Gervais  and  Phalen.  In 
order  to  get  sufficient  pressure  to  supply  the  higher  areas,  the 
water  is  forced  into  a  reservoir  one  mile  west  of  the  pumping 
station.  This  reservoir  is  290  feet  above  the  water  level  of  the 
Mississippi  river  and  has  a  capacity  of  18,000,000  gallons. 
There  is  another  reservoir  on  the  West  Side  to  supply  the  ele- 
vated district  across  the  river. 

At  the  pumping  station  and  also  at  the  entrance  of  the  con- 
duit leading  from  Lake  Vadnais,  a  series  of  graduated  wire 
screens  strain  from  the  water  the  coarser  vegetable  growth. 

609 


610  MINNESOTA    BOTANICAL    STUDIES. 

Method  of  collection. — The  method  of  collection  is  practically 
the  one  suggested  by  Dr.  Smith  Ely  Jelliffe  *  and  is  as  follows  : 
A  piece  of  absorbent  cotton  four  or  five  inches  square  and  one 
inch  thick  is  attached  by  means  of  a  twelve-inch  square  of  un- 
bleached muslin  to  the  water  faucet.  The  water  is  then  turned 
on  sufficiently  to  insure  a  constant  stream  and  is  allowed  to  run 
from  ten  to  twelve  hours,  after  which  the  cotton  is  removed. 
The  cotton,  which  is  usually  quite  brown  from  the  organisms, 
is  divided  into  pieces  and  rubbed  and  rinsed  in  five  beakers  each 
containing  200  c.c.  of  water.  The  water  is  then  poured  into 
one  vessel  and  allowed  to  settle,  after  which  the  deposit  is  put 
into  a  glass  containing  25  c.c.  A  few  drops  of  this  is  trans- 
ferred to  a  slide  by  means  of  a  pipette  and  examined  microscop- 
ically. At  least  ten  mounts  from  each  week's  collection  of  ma- 
terial was  examined  in  this  wa}^.  The  rest  was  then  preserved 
in  2  per  cent,  formaline  for  future  reference. 

For  the  records,  Dr.  Jelliffe's  method  of  computation  was 
adopted.  In  computing  the  numbers  the  following  schedule 
was  used  : 

Abundant,  25  +  in  one  c.c.  of  water. 

Common,  10-25  in  one  c.c.  of  water. 

Few,  5-10  in  one  c.c.  of  water. 

Scarce,  1-5  in  one  c.c.  of  water. 

Present,  Less  than  five  in  one  c.c.  of  water. 

Since  November,  1899,  weekly  collections  have  been  made  of 
the  plant  life  in  the  St.  Paul  water  supply  and  the  organisms 
identified  (as  far  as  possible)  and  their  number  computed. 

The  vegetable  organisms  found  were  all  algas  if  we  except 
the  pollen  grains  and  Fungi  spores  that  appeared  occasionally. 
The  Algse  found  were  as  follows  : 

1.  Diatomacece,  13  varieties. 

2.  Cyanophycece,  n  varieties. 

3.  Cklor<rphyce<&)  32  varieties. 

4.  PeridinicR,  2  varieties. 

Of  these  some  forms  of  Diatoms  were  present  almost  con- 
stantly, especially  Melosira,  Stephanodiscns,  two  varieties  of 

*  Jelliffe,  S.  E.  A  preliminary  report  upon  the  microscopical  organisms  found 
in  the  Brooklyn  water  supply.  Brooklyn  Med.  Journ.  7 :  595.  O.  1893. 


Panning:    ALG^E  OF  THE  ST.  PAUL  CITY  WATER.  611 

Synedra  and  Asterionella.  Of  the  Chlorophyceae,  Scenedes- 
mus  and  Raphidium  were  present  most  of  the  time.  Among 
the  Cyanophycere,  Oscillatoria  and  Ccelospharium  were  prac- 
tically constant. 

The  effect  of  the  cold  on  some  of  the  varieties  is  shown  by 
the  accompanying  plates  (PI.  XLIII.  and  XLIV.).  It  will  be 
seen  that  a  fall  in  temperature  coincides  with  a  decrease  in  the 
numbers  of  all  except  Oscillatoria,  which  shows  a  gradual  in- 
crease with  the  cold  and  is  abundant  for  several  weeks  during 
the  severest  weather. 

The  desmids,  more  plentiful  in  the  fall  than  at  any  other 
time,  although  never  abundant,  disappeared  after  the  cold  wave 
of  the  third  week  in  December.  Another  fall  in  temperature 
about  the  fourth  week  in  January  banished  most  of  the  Chloro- 
phyceae and  cleared  the  field  of  Ccelosphcerium,  but  some  of  the 
diatoms  persisted  until  the  zero  weather  in  February,  when 
most  of  them  disappeared.  Fragilaria  was  abundant  in  Jan- 
uary and  about  January  3Oth,  when  Oscillatoria  and  Fragilaria 
were  practically  the  only  forms  seen,  the  water  contained  con- 
siderable sandy  debris  and  entangled  in  it  were  quantities  of 
resting  spores  of  Glceotrichia. 

The  observations  vary  from  year  to  year  so  that  a  record 
should  be  kept  for  several  years  before  one  could  find  what 
forms  were  both  constant  and  abundant.  For  example,  in  the 
fall  of  1898,  Anabcena  was  "  common,"  but  in  the  following 
year  it  is  only  marked  "  present." 

I  wish  to  thank  Mr.  P.  F.  Lyons,  of  the  St.  Paul  Weather 
Bureau  and  Mr.  John  Caulfield,  Secretary  of  the  Water  Board, 
St.  Paul,  for  their  kind  assistance. 

EXPLANATION  OF  PLATES  XLIII.  AND  XLIV. 
Table  showing  relation  between  abundance  of  certain  forms  and 
temperature.     XLIV.  is  a  continuation  of  XLIII. 

PLATE  XLV. 

1 .  Coelastrum  microporum  Naeg.  var.  speciosum  Wolle.  Freshw. 
Algae  U.  S.  170.  pi.  156.  f.  4.     1887. 

2.  Pediastrum  duplex  Meyen  Beob.  iiber  Algenformen.  in  Nova 
Acta  Acad.  Leop.  Carol.  772.      1829. 

3.  Scenedesmus  quadricauda  (Turp.)  Breb.  Alg.  Falais.  66.    1835. 

4.  Scenedesmus  bijttgatus  (Turp.)  Kg.  Syn.  Diat.  607.      1833. 

5.  Rhaphidium  polymorphum   Fresen.  var.    aciculare    (A.   Br.) 
Rabenh.  Fl.  Eur.  Algar.  3:  45.     1868. 


612  MINNESOTA    BOTANICAL    STUDIES. 

6.  Rhaphidium    polymorphism   Fresen.    var.   falcatum    (Corda) 
Rabenh.    Fl.  Eur.  Algar.  3:  45.      1868. 

7.  Nephrocytium  agardhianum  Naeg.  Gatt.  einz.  Alg.  So.  pi.  3  C. 
1849. 

8.  Dictyosphcerium  pulchellum  Wood.  Freshw.  Algse  U.  S.  84. 
1873- 

9.  Eridorina  elegans  Ehrenb.  in  Monatsb.  der  Akad.  d.  wiss.  zu 
Berlin.   78,  152.  pi.  2.f.  10.      1831. 

10.  Pandorina  morum  (Muell.  ?)  Bory  in  Ehrenb.  Infus.  53.  pi. 
2.f.33-     1838. 

11.  Glozocystis  gigas  (Kg.)   Lagerh.  Bidrag.  till  Sveriges  Algfl. 
63.      1883.    ' 

12.  Micrasterias  truncata   (Corda)  Bre"b.  in  Ralfs  Brit.  Desmid. 
75.  no.  9.  pi.  8.  f.  4.  and  pi.  10.  f.  5.     1848. 

13.  Staurastrum  sebaldi  Reinsch.  Algenf.  von  Franken.  175.  pi. 
n.  f.  i.      1867. 

14.  Staurastrum  minneapoliense  Wolle  in  Bull.  Torr.  Bot.  Club, 
12:  5.  j07.  47.  f.  11-13.      1885. 

15.  Staurastrum     paradoxum     Meyen.     var.     longipes    Nordst. 
Sydlig.  Norg.  Desm.  35. /.  17.     1873. 

1 6.  Arthrodesmus  incrassatus    Lagerh.   var.   cycladatus   Lagerh. 
Bidrag  till  Amer.  Desm. -Flora.  242.  pi.  27.  f.  19.      1885. 

17.  Pleurotceniopsis  quaternaria  (Nordst.)  De  Toni.  Syll.  Algar. 
I :  914.      1889. 

18.  Cosmarium  nitidulum  De  Not.   Element.   42.   pi.  3.  f.   26. 
1867. 

19.  Closter.ium  parvulum  Naeg.  Gatt.  einz.  Alg.  106.  pi.  6.   C.  f. 

2.       1849. 

20.  Nostoc  sp.  und. 

21.  Anabcena  Jlos-aqu<z    (Lyngb.)  Breb.    Algues  des  environs  de 
Falaise  36.      1835. 

22.  Lyngbya    majuscula    Harv.    in    Hooker,    Eng.    Fl.    5:    370. 
1833- 

23.  Merismopedia  g-faucaNaeg.  Gatt.  einz.  Alg.  55.  pi.  i.  D.f. 
I.      1849. 

24.  Ccelosphceriiim  kutzingianum  Naeg.  Gatt.    einz.  Alg.  54.  pi. 
i.  C.     1849. 


Fanning:   ALG^E  OF  THE  ST.  PAUL  CITY  WATER.  613 

25.  Anacystis  marginata  Menegh.  Consp.  6.      1837. 

26.  Peridinium  tabulatnm  Ehr.  in  Kent.  Manual  of  the  Infusoria. 
I :   448. 

PLATE  XLVI. 

1.  Amphora  ovalis  (Brdb.)  Kg.  Bac.  107.  pi.  5.  f.  jj",  jp.  1844. 

2.  Cymbella  lanceolata   (Ehr.)  Kirchn.  Alg.   Schles.  iSS.  1878. 

3.  Asterionella  formosa  Hass.  in  Micr.  Exam.  10. 

4.  Synedra  ulna  (Nitzsch.)  Ehr.  Infus.   21 1.  //.  77.  f,  i.  1836. 

5.  Synedra  pulchella  (Ralfs.)  Kg.  Bacill.  68.  pi.  29.  f.  37.  1844. 

6.  Fragilaria  capiicina  Desmaz.  Crypt,    de  France  (eel.    i),  no. 
453-     1825. 

7.  Tabellaria  fenestrata  Lyngb.  Kg.  var.  intermedia  Griin.    in., 
V.  H.  Syn.  //.  52.  f.  6-8.     1 880-81. 

8.  Cyclotella  comta  (Ehr.)  Kg.  Spec.  Algar.  20.      1849. 

9.  Stephanodiscus  niagarac  Ehr.  in  Berl.  Akatl.  So.      1845. 

10.  Melosira  granulata  (Ehr.)  Ralfs.  in  Pritch.  Inf.  820.      ^845- 
61. 

1 1 .  Ceratium  longicorne  Carter. 


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VOL.  II. 


MINNESOTA  B( 


Oct. 


Nov.      Dec.       Jan.        Feb. 


Abund. 

Temperature    £•„ 

\ 

ol) 

Common       7c° 

\ 

/  \J 

~in° 

\ 

/u 
Few             oc. 

\ 

DO 

Melosira             gg» 

\ 

Scarce        ci;» 

\ 

cn« 

\ 

DU 

Present        A*;0 

\ 

1 

T"U 

An° 

\ 

^ 

—— 

X 

V 

1 

Absent        -jt;0 

s 

\ 

/ 

Ov 
•3  no 

\ 

JU 

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\ 

t 

/ 

9(1° 

"X 

r 

\ 

\l 

\ 

/ 

Common         ic° 

\ 

3 

\ 

/ 

i  j 
in0 

\ 

/ 

\i 

V 

/ 

lu 
Few                 co 

\ 

\\ 

/ 

\ 

1 

J 

Synedra                n° 

/ 

\ 

r 

Scarce 

\ 

\ 

Present 

\ 

\ 

/ 

\ 

/ 

Absent 

\ 

/ 

\ 

/ 

Abund. 

\ 

/ 

V 

/ 

\ 

Common 

\ 

/ 

3 

/ 

\ 

\ 

/ 

\ 

Few 

\ 

/ 

^ 

Stephanodiscus 

\ 

Scarce 

\ 

\ 

/ 

Present 

\ 

/ 

Absent 

PLA 


THE    HELIOTVP 


;iCAL  STUDIES. 


PART  V. 


Apr         May       June     July          Aug.      Sept. 


VOL.  II. 


MINNESOTA  BC 


Oct.        Nov.        Dec.  Jan.          Feb.         f 


Abund. 

Temperature  F. 
fifl" 

/ 

\ 

Comrnon      75° 

/ 

\ 

70° 

\ 

I 

Few             RK° 

\ 

f 

Coelosphaerium   50° 

\ 

/ 

\ 

\ 

\ 

Sc3trc6         R^* 

\ 

/ 

\ 

I 

/ 

'in0 

\ 

^* 

\ 

/ 

\ 

/ 

Present       45° 

\ 

1 

\ 

/ 

Af\" 

\ 

,  — 

,  —  • 

-\ 

\ 

/ 

Absent        35° 

\ 

\ 

f 

?n« 

\ 

Abund.         ?v> 

\ 

/ 

/ 

\ 

?ne 

/ 

\ 

/ 

1 

\ 

/ 

Common       15* 

/ 

\ 

/ 

^ 

/ 

in° 

\ 

/ 

\\ 

i 

S 

Few               5° 

\\ 

/ 

\ 

/ 

Scenedesmus         0° 

\ 

1 

V 

/ 

V 

j 

7 

Scarce 

\ 

1 

\ 

/ 

\ 

I 

Present 

\ 

Absent 

Abund. 

/ 

!\ 

/ 

\ 

Common 

/ 

\ 

/ 

\ 

/ 

Few 

r  • 

Oscillatoria. 

/ 

Scarce 

/ 

\ 

/ 

Present 

\ 

/ 

Absent 

IJpAL  STUDIES. 


PART  V. 


A  r.          May          June        July         Aug.  Sept 


K 


\ 


\ 


:LIV. 


Q   CO.,    BOSTON. 


VOL.  II. 


MINNESOTA 


CAL  STUDIES. 


PART  V. 


11 


10 


23 


obgooo 


ooOBoBSB 


22 


VOL.  II. 


MINNESOTA  BOTANICAL  STUDIES. 


PART  V. 


PLATE  XLVI. 


THE    MELIOTVP€    PUINTIKC   CO.,    BOSTON. 


XXXV.    NOTES    ON   SOME   PLANTS    OF   ISLE 
ROYALE. 


W.  A.  WHEELER. 


During  August,  1900,  I  spent  about  two  weeks  at  Tobin  Har- 
bor on  the  eastern  end  of  Isle  Royale,  Mich.  At  this  time  I 
made  a  collection  of  plants  numbering  about  150  species,  of 
which  the  following  seem  to  be  worthy  of  note. 

Botrychium  lunaria  (L.)  Sw.  Schrad.  Journ.  Bot.  2  :  no.     1800. 
Rare  in  moist  woods  and  thickets. 

Botrychium  virginianum  (L.)  Sw.  Schrad.  Journ.  Bot.  2  :   in. 

1800. 

Prothallia  and  young  sporophytes  were  collected  in  a  thicket 
near  the  shore  of  the  lake  on  Aug.  30.  Previous  collections  of 
the  prothallia  of  this  species  have  been  made  only  by  Professor 
Douglas  H.  Campbell  at  Grosse  Isle,  Michigan,  in  1893  and  by 
Professor  E.  C.  Jeffrey  at  Little  Metis,  Quebec,  Canada,  in 
1895. 

Woodsia  ilvensis  (L.)  R.  BR.  Trans.  Linn.  Soc.  n  :   173.     1812. 
Very  common  on  exposed  rocks  along  the  lake  shore. 

Dryopteris  fragrans  (L.)  SCHOTJ,  Gen.  Fil.     1834. 
Common  on  exposed  rocks  with   Woodsia  ilvensis. 

Cryptogramme  acrostichoides  R.   BR.   App.  Franklin's  Journ. 

767.     1823. 

Infrequent  on  rocks.  This  is  probably  one  of  the  rarest  ferns 
of  the  Great  Lake  region. 

Lycopodium  selago  L.  Sp.  PL  1102.     1753. 
Frequent  on  rocks  near  the  shore. 

Selaginella  selaginoides  (L.)  LINK,  Fil.  Hort.  Berol.  158.     1841. 
Frequent  on  moist,  shaded  rocks  near  the  water's  edge  on  the 
shore  of  the  harbor. 

619 


620  MINNESOTA    BOTANICAL    STUDIES. 

Calamagrostis  langsdorfii  (LINK.)  TRIN.  Unifl.  225.     1824. 

Previously  collected  from  Isle  Royale  in  1865  by  T.  C. 
Porter.  This  may  be  the  collection  reported  in  Beal  & 
Wheeler's  Flora  of  Michigan  as  C.  lapponica  Trin. 

Carex  abacta  BAILEY,  Bull.  Torr.  Club,  20:  427.     1893. 
Carex  limosa  L.  Sp.  PL  977.     1753. 
Juncus  articulatus  L.  Sp.  PL  327.     1753. 

Sagina  nodosa  (L.)  FENZL,  Verbr.  Alsin.  18.     1833. 
Infrequent  on  moist  rocks. 

Sisymbrium  humile  MEYER,  in  Ledeb.  Fl.  Alt.  3:    137.     1831. 
Rare  on  rocks. 

Echinopanax  horridum  (SMITH)  DEC.  &  PLANCH.  Rev.  Hortic. 

105.     1854. 

The  occurrence  of  this  plant  in  a  locality  so  far  removed  from 
what  had  been  considered  its  native  home  is  certainly  remark- 
able. No  collections  of  it  have  been  reported  farther  east  than 
the  Rocky  Mountains  of  Montana  and  British  Columbia.  On 
Isle  Royale  it  occurs  on  the  rocky  cliffs  at  two  places  near 
Tobin's  Harbor,  i.  <?.,  Passage  Island  and  Black's  Point  and 
has  the  appearance  of  being  indigenous. 

Gentiana  rubricaulis  SCHWEIN,  in  Keating's  Narr.  Long's  Exp. 

2  :  384.      1824. 
Frequent  in  moist  open  places  and  on  protected  rocks. 

Castilleja  acuminata  (PURSH)  SPRENG.  Syst.  2:  775.     1825. 
Euphrasia  americana  WETTST.  Mon.  Euph.  127.     1896. 
Aster  lindleyanus  T.  &  G.  Fl.  N.  A.  2  :  122.     1841. 

Erigeron  acris  droebachianus  (O.  F.  MUELLER)  BLYTT.  Norg. 
Fl.  i:  562.      1861. 

Senecio  discoideus  (HooK.)  BRITTON,  in  Britton  &  Brown,  111. 
Fl.  3:  479.     i! 


XXXVI.     REVEGETATION    OF    TRESTLE    ISLAND. 


D.  LANGE. 


The  piece  of  land  I  have  named  Trestle  island  lies  in  the 
southwest  part  of  Lake  Phalen,  near  St.  Paul,  Minnesota.  The 
island  consists  of  an  East  and  a  West  section  separated  from 
each  other  by  a  twenty-five  foot  embankment  of  the  St.  Paul 
&  Duluth  Railroad. 

THE  SEASON  OF  1898. 

Up  to  the  spring  of  1898,  the  road  maintained  a  trestle  over 
the  shallow  southeast  bay' of  Lake  Phalen  in  place  of  the  present 
embankment.  When  the  trestle  was  filled  in  between  April 
10,  and  June  i,  1898,  the  gradually  increasing  weight  of  the 
dumped  material  caused  the  soft  lake  bottom  marl  which,  ac- 
cording to  the  statement  of  the  road's  engineer,  is  from  6  to  35 
feet  thick,  to  slip  out  laterally  ;  and  with  many  folds,  wrinkles 
and  fissures  it  rose  from  a  few  inches  to  ten  feet  above  low  water 
level  of  the  lake.  Although  the  engineer  in  charge  tried  to 
prevent  the  slipping  of  the  marl  by  means  of  pontoons,  the 
movement  did  not  cease  until  about  2500  square  yards  of  lake 
bottom  had  risen  and  become  dry  land.  Of  this  land  about 
1500  square  yards  rose  east  of  the  track  out  of  6  inches  to  3 
feet  of  water,  and  will  be  called  the  East  section  in  this  paper, 
the  other  1000  square  yards  rose  West  of  the  track  out  of  3  to  5 
feet  of  water  and  will  be  referred  to  as  the  West  section.  Both 
sections  have  been  under  the  writer's  observation  from  April 
1898,  to  October  15,  1900,  and  it  is  intended  to  show  in  this 
paper  the  most  marked  changes  in  the  vegetation  of  Trestle 
island. 

Early  in  June,  1898,  both  sections  presented  a  curious  system 
of  curved  ridges  and  crevices  running  generally  parallel  to  one 
another  and  looking  like  miniature  mountain  ridges,  valleys, 
and  gaping  faults  ;  and  even  small  lakes  with  snails  and  other 
aquatic  creatures  were  to  be  seen.  Many  of  the  crevices  were 

621 


622  MINNESOTA    BOTANICAL    STUDIES. 

over  two  feet  deep  and  while  the  opposite  walls  approached  each 
other  below,  the  gap  at  the  surface  was  in  many  cases  eight  inches 
wide.  The  West  section  was  an  expanse  of  mud  ridges  entirely 
bare  of  vegetation,  and  was  at  first  too  soft  to  admit  of  a  man 
passing  over  it ;  the  East  section,  however,  having  risen  out  of 
shallower  water  and  partly  out  of  a  marsh,  exhibited  specimens 
of  yellow  pond  lilies  (Nymphea  advena),  cattails,  water  plan- 
tain, and  common  rush  lifted  out  of  the  water  and  struggling 
under  adverse  conditions. 

In  the  early  part  of  July,  1898,  the  East  section  looked  already 
green  from  the  distance.  The  aquatic  plants  just  mentioned 
still  lived,  but  showed  the  effects  of  changed  conditions.  The 
leaves  of  Nymphea  advena,  for  instance,  were  all  very  short- 
petioled,  and  were  below  normal  size,  appeared  more  or  less 
brownish  and  the  younger  ones  were  rolled  from  both  margins 
inward  and  upward.  On  the  other  hand,  the  number  of  true 
land  plants  growing  vigorously  was  already  bewildering  and 
although  most  of  them  were  still  too  young  for  a  reliable  inden- 
tification,  the  following  were  found  in  bloom  about  July  5,  1898  ;* 

1.  White  clover  {Trifolium  r  opens]. 

2.  Red  clover  (Trtfolmm  pratense). 

3.  Wild  mustard  (Brassica  nigra]. 

4.  Peppergrass  sp. 

5.  Mayweed  (Anthemis  cottild]. 

6.  Crucifer  sp. 

7.  Black  nightshade  (Solatium  nigruni). 

8.  Mustard  sp. 

9.  Polygonum  sp. 

10.  Timothy  grass  (Phleum  pratense]. 

11.  Grass  sp. 

12.  Kentucky  bluegrass  {Poa  pratensis),  out  of  bloom. 

The  southern  part  of  this  section  was  covered  with  a  compact 
layer  of  fine  silt  from  the  railroad  embankment  and  on  this  firm 
soil  little  else  but  young  mosses  were  growing.  Scattered  over 
the  higher  part  of  the  whole  section  were  young  cottonwoods 
and  willows. 

The  most  interesting  plants  found,  however,  were  the  pro- 
thallia  of  horsetails,  probably  of  Equisehim  arvense,  as  that 


*  The  botanical  nomenclature  of  this  paper  follows  the  "  Illustrated  Flora  "  of 
Britton  and  Brown. 


Lange  :    REVEGETATION  OF  TRESTLE  ISLAND.  623 

species  is  very  common  along  the  railroad  track  south  of  the 
island.  These  prothallia  were  little  green  clumps  from  */£  to 
y^  of  an  inch  in  diameter  and  grew  mostly  on  the  walls  of 
the  shaded  mud  cracks.  From  many  of  them  one  or  more 
thin  young  horsetails  were  protruding.  Although  I  carefully 
searched  the  cracks  for  these  prothallia  in  the  springs  of  1899 
and  1900,  I  did  not  again  find  a  single  specimen.  In  1898 
they  were  abundant  on  both  sections  of  the  island,  but  they  did 
not  grow  on  the  marl,  but  only  on  silt  and  in  silt  cracks.  This 
silt  consisted  of  a  brownish  clay  and  of  very  fine  quartz  sand, 
making  a  compact  damp  soil. 

The  West  section  was  still  quite  bare.early  in  July,  1898.  The 
creviced  marl  ridges  had  dried  in  the  sun  and  exhibited  white 
streaks  like  limestone.  These  little  crags  and  points  were  bare, 
but  in  shaded  and  sheltered  depressions  mosses  covered  much 
of  the  damp  marl ;  and  small  cottonwoods,  aspens  and  willows 
were  scattered  over  the  whole  West  section. 

About  September  loth  the  aspect  of  both  sections  had 
changed,  but  most  markedly  that  of  the  East  section,  which 
was  a  veritable  wilderness  of  weeds.  Its  lower  portion  was 
covered  with  wild  rice,  over  six  feet  high.  Under  the  rice  and 
also  on  the  higher  ground  water  hoarhound  (Lycopus  ameri- 
canus],  skullcap  (Scutellaria  laterifolia)  and  American  wild 
mint  (Mentha  canadensis]  formed  dense  tangles  and  grew  with 
a  luxuriance  I  had  never  seen  before.  There  were  present  in 
great  confusion  nearly  all  the  plants  mentioned  in  the  count  of 
August  16,  1899,  and  a  f£W  not  found  at  that  day.  In  this 
weeds'  paradise  the  young  willows  could  be  seen,  but  they  were 
not  at  all  conspicuous.  No  bare  ground  was  now  visible  on 
this  section,  but  in  walking  through  the  weeds,  one  could  not 
avoid  frequent  stumbling  into  the  cracks.  The  West  section  at 
this  time  presented  the  appearance  of  a  loose  growth  of  young 
cottonwoods,  aspens  and  willows,  with  the  cottonwoods  most  and 
the  willows  least  conspicuous,  but  much  bare  soil  could  still  be 
seen  even  from  a  distance. 

On  both  sections  all  the  young  trees  remained  green  and  con- 
tinued to  grow  until  late  in  the  fall.  On  October  22d  they  were 
still  green  but  had  evidently  ceased  growing  on  account  of  the 
cool  weather  that  had  prevailed  since  the  third  of  the  month. 
Of  the  East  section  my  notes  for  October  22d  say:  "A  great 
wilderness  of  dead  weeds." 


624  MINNESOTA    BOTANICAL    STUDIES. 

THE  SEASON  OF  1899. 

Although  Trestle  island  was  visited  during  the  winter  of 
1898-99  and  was  also  observed  during  the  spring  and  early 
summer  of  1899,  I  shall  at  once  proceed  to  a  midsummer  sketch 
of  it,  as  it  appeared  from  August  14  to  16,  1899. 

Compared  with  the  preceding  season,  the  West  section  had 
this  time  changed  most  rapidly  and  radically.  The  mud  flat  of 
fourteen  months  ago  was  now  a  thicket  of  young  peach-leaved 
willows  (Salix  amygdaloides).  The  cotton  woods  (Populus 
deltoides)  and  the  aspens  (Populus  tremul aides]  so  conspicu- 
ous last  year  were  now  hidden  by  the  willows,  which  had  grown 
so  large  that  a  herd  of  cattle  or  horses  would  have  been  com- 
pletely concealed.  Although  the  high  railroad  embankment 
was  not  more  than  forty  feet  from  the  spot  where  I  wrote  up 
my  notes,  I  could  only  get  glimpses  of  it  along  the  sky  line. 
Within  fourteen  months  there  had  grown  from  the  tiny,  wind- 
carried  willow  seeds  a  thicket  of  trees  that  were  large  enough 
to  completely  shade  my  paper  and  exclude  the  southwest 
breeze,  while  two  of  the  young  trees  were  stout  enough  to 
afford  me  a  secure  and  comfortable  back  rest.  One  of  them, 
by  actual  measurement,  was  n  feet  3  inches  high  and  had  a 
diameter  of  i^  inches  at  a  distance  of  four  inches  above  the 
ground.  All  the  larger  trees  of  this  species  were  approxi- 
mately of  that  size  and  showed  a  growth  of  about  five  feet  for 
the  season.  A  few  of  the  cottonwoods  not  standing  very  close 
to  any  Salix  amygdaloides  were  about  as  tall,  but  not  as  thick 
as  the  Salix,  while  the  aspens  and  three  or  four  other  species  of 
willow  were  much  smaller  and  showed  only  a  growth  of  about 
three  feet  for  the  season.  It  was  clear  that  the  peach-leaved 
willow  would  be  the  dominant  plant  on  this  section.  It  has 
clearly  won  the  battle  against  the  cottonwoods  ;  and  other  trees 
and  herbs  will  occupy  a  subordinate  position.  The  West  sec- 
tion is  a  willow  island. 

Lack  of  space  forbids  to  enlarge  upon  the  herbs  and  grasses 
on  this  section,  but  it  should  be  mentioned  that  a  fringe  of  cat- 
tails {Typha  latifolia]  and  arrowheads  (Sagittaria  latifolia)  is 
forming  on  the  lake  side  of  the  West  section.  These  plants 
grow  there  in  6  to  12  inches  of  water,  where  the  whitish  marl 
was  not  raised  above  low  water  level  although  it  was  elevated 
considerably. 


Lange:    REVEGETATION  OF  TRESTLE  ISLAND.  625 

The  East  Section  from  August  14  to  16,  1899. — The  great  di- 
versity of  plant  life  on  this  section  already  referred  to  continued 
for  this  season.  In  this  diversity  and  in  the  comparative  pau- 
city of  trees  it  presented  a  striking  contrast  to  the  West  section, 
where  all  true  land  herbs  struggled  under  the  over-shadowing 
trees. 

On  the  East  section,  peach-leaved  willows  dominated  the 
south  corner.  A  crescent,  which  under  the  effect  of  higher 
water,  formed  last  year  a  dense  rice  marsh,  with  a  thick  tan- 
gled undergrowth  of  water  hoarhound  and  mad-dog  skullcap 
is  now  a  meadow  of  tangled  rice  cut-grass  (ffomolocenchrus 
oryzoidcs].  The  remainder  of  the  section  is  covered  by  a  very 
much  mixed  vegetation,  amongst  which  peach-leaved  willows, 
slender  pink  persicaria,  giant  sunflower,  a  few  specimens  of 
wild  rice,  and  cattail  are  most  conspicuous  although  not  most 
numerous. 

The  most  notable  changes,  besides  the  one  of  the  rice  marsh 
to  a  cut-grass  meadow,  are  the  complete  disappearance  of 
water-lilies,  wild  rice,  and  common  rush  from  the  higher  ground, 
the  appearance  of  slender  pink  persicaria  on  well-marked  areas, 
and  the  establishment  of  willow  domination  in  the  south  corner. 

Not  a  hand's  breadth  of  bare  soil  is  any  longer  visible  on 
either  section,  except  in  deep  crevices.  To  show  clearly  the 
distribution  of  plant  life  on  this  section  in  midsummer  of  1899, 
I  give  here  the  result  of  a  count  made  on  August  16,  1899. 

I.  Plants  established  in  large  numbers  and  forming  the  bulk 
of  the  vegetation. 

1.  Peach-leaved  willow  (Salix  amygdaloides).     In  the  south 
corner. 

2.  Rice  cut-grass  (ffomolocenchrus  oryzoides).     On  an  east 
crescent. 

3.  Slender  pink   persicaria   (Polygonum   incur  natuni).     In 
well-defined  patches. 

4.  Mad-dog    skullcap    (Scutellaria    later  if  olio).      General 
under  the  taller  plants. 

5.  Cut-leaved    water    hoarhound     (Lycopus    americanus). 
General  under  taller  plants. 

6.  American  wild  mint  (Mentha  canademis).    General  under 
taller  plants. 

7.  Common  ragweed  (Ambrosia  artemisi&folia}.      On  high- 
est ridges,  where  common  rush  grew  in  '98. 


626  MINNESOTA    BOTANICAL    STUDIES. 

8.   Horseweed  (Leptilon  canadense).     On  the  higher  ridges, 
where  common  rush  grew  in  '98. 

II.  Aquatics,  marsh  plants  and  cryptogams  which  were  still 
struggling  along  in  a  few  spots.    July  of  this  year,  the  most  im- 
portant month  of  growth  for  annual  and  perennial  herbs  in  this 
region   was   unusually  dry,   showing  a  monthly   deficiency  in 
precipitation  of  1.75  inches.*     This  condition  was  unfavorable 
for  aquatics  and  marsh  plants,  but  favorable  for  such  plants  as 
horseweeds  and  common  ragweeds. 

1.  Wild  rice  (Zizania  aquaticd). 

2.  Water  plantain  (Altsma  -plantago-aqiiatica). 

3.  Common  rush  {Juncus  cffusns}. 

4.  Broad-leaved  cattail  (Typha  lalifolia). 

5.  Tuberous   white  water  lily  (Castalia  tuberosa).     Identi- 
fied from  leaves  only. 

6.  Large    yellow    pondlily    {Nymphea    advena}.       Leaves 
only,  a  flower  bud  found  an  inch  deep  in  wet  soil. 

7 .  Horsetails  ;  mosses,  amongst  them  Funaria  hygrometrica  ; 
liverworts,  with  Marchantia  polymorpha  fairly  common. 

III.  Plants  of  which  only  a  few  scattered  specimens  were 
found. 

1.  Slender  nettle  (Urtica  gracilis], 

2.  Swamp  milkweed  (Ascleptas  incarnatd). 

3.  Smooth  bur-marigold  (Bidens  lavis}. 

4.  Marsh  skullcap  (Scutettaria  galericulata). 

5.  Blue  vervain  (  Verbena  hastatd]. 

6.  White  vervain  (  Verbena  urticifolid). 
7-  Beggarticks  (Bidens  frondosa}. 

8.  Aster-like  boltonia  (Boltonia  asteroides}. 

9.  Rough  cinquefoil  (Potentilla  monspeliensis). 

10.  Great  ragweed  (Ambrosia  trifida}. 

11.  American  cocklebur  (Xanthium  canadense]. 

12.  Prickly  lettuce  (Lactuca  scariola). 

13.  Lactuca  sp. 

14.  Heart-leaved  willow  (Salix  cordata], 

15.  Salix  sp. 

16.  Cottonwood  (Popuhis  deltoides], 

17.  American  aspen  (Populus  tremuloides}. 

18.  Sullivant's    milkweed  (Asclepias  sullivantii}.     Identified 
from  leaves  only. 

*The  rainfall  data  of  this  paper  are  taken  from  the  Monthly  Meteorological 
Summary  of  the  United  States  Weather  Bureau  for  St.  Paul,  Minn. 


Lange:  REVEGETATION  OF  TRESTLE  ISLAND.  627 

19.  Water  pepper  (Polygonum  hydropiper). 

20.  Climbing  false  buckwheat  {Polygonum  scandens). 

21.  Lamb's  quarter  (Chenopodinm  sp.). 

22.  Dock  (Rtimex  sp.). 

23.  Giant  sunflower  (Helianthns  giganteus). 
i\.  Black-eyed  Susan  {Rndbcckia  htrta). 

25.  Common  thoroughwort  (Eupatorhim  pcrfo liatu m). 

26.  Late  goldenrod  (Solidago  seroti.no). 

27.  Canada  goldenrod  {Solidago  canadensis}. 

28.  Prairie  mugwort  {Artemisia  gnaphaloides}. 

29.  Bushy  aster  (Aster  dumosus}.    Identified  from  leaves  only. 

30.  Red  clover  (Trifolium  pratcnse], 

31.  Peppergrass  (I^epidium  sp.). 

32.  Tall  sisymbrium  (Sisymbrium  altissimutri). 

33.  Black  mustard  (Brassica  nigrd]. 

34.  Black  nightshade  (Solanum  nigruni). 

35.  Bristly  buttercup  (Ranunculus  pennsylvanicus). 

36.  Swamp  willow-herb  (Epilobium  palustre). 

37.  Common  evening  primrose  (Onagra  biennis). 

38.  Canada  thistle  (Carduus  arvensis). 

39.  Common  thistle  (Cardutts  Janceolattis}. 

40.  Water  hemlock  (Cictita  maculata). 

41.  Nodding  wild  rye  (Elymus  canadensis]. 

42.  Squirrel-tail  grass  (Hordeum  jubatum}. 

43.  Grass  sp. 

44.  Grass  sp. 

45.  Grass  sp. 

46.  Grass  sp. 

47.  Sedge  (Cyperus  sp.). 

THE  SEASON  OF  1900. 

The  last  half  of  October,  1899,  was  <luite  ^T  and  at  the  end 
of  the  month  the  United  States  Weather  Bureau  reported  an 
accumulated  deficiency  in  precipitation  since  January  first  of  .73 
inch.  On  some  day  between  October  29  and  November  4, 
1899,  a  prominent  factor  in  plant  distribution  appeared  on  the 
East  section  of  Trestle  island.  Fire  changed  the  wilderness 
of  dead,  dry  weeds  into  a  black,  ashy  waste. 

On  May  6,  1900,  the  East  section  looked,  on  the  whole,  still 
barren  and  black.  All  the  over-ground  parts  of  the  young 


028  MINNESOTA    BOTANICAL    STUDIES. 

trees  that  had  grown  scattered  amongst  the  weeds  were  found 
to  have  been  fire-killed,  but  about  one-half  of  their  number  had 
grown  out  again  near  the  ground.  The  larger  ones  in  the  south 
corner  had  not  been  killed,  because  there  the  weeds  had  not 
(jrown  as  thick. 

O 

Patches  of  the  perennials,  Elytnus  canadensis  and  a  few  other 
grasses,  as  well  as  goldenrods,  sunflowers  and  nettle  were  coming 
out  vigorously.  The  rice  cut-grass,  however,  although  it  seems 
to  be  perennial,  had  a  very  poor  start.  On  close  examination 
numerous  seedlings  of  annuals  were  found,  but  all  looked 
brownish  and  sickly  on  account  of  the  dry  weather  and  the 
glaring  sunlight.  Although  most  of  them  were  not  readily 
identified,  I  recognized  the  following  without  difficulty : 

1.  Polygonum  sp. 

2.  Mentha  sp. 

3.  Carduus  lanceolatus,  the  seedlings  of  which  looked  quite 
green  and  vigorous. 

4.  Lactuca  sp. 

5.  Brassica  nigra. 

6.  Ckenopodium  sp. 

The  spring  and  summer  of  1899  were  excessively  dry  in  this 
region  so  that,  since  January  ist  a  continued  accumulated  defi- 
ciency of  rain  of  over  4  inches  was  reported  from  May  3ist 
to  August  3ist.  About  June  loth  another  new  factor,  one  that 
is  able  to  destroy  all  plant  life  not  perennial  or  not  armed  in 
some  way,  appeared  on  the  scene — cattle,  a  large  herd  of 
hungry  dairy  cows.  In  their  search  for  green  grass  they  had 
passed  over  the  dried-up  marsh  and  along  at  the  foot  of  the 
embankment  and  had  discovered  the  East  section  of  Trestle 
island. 

In  the  end,  however,  bovine  instinct  for  plant  selection  proved 
of  some  interest,  for,  when  after  an  absence  of  over  two  months, 
I  visited  the  place  again  about  Sept.  loth,  there  was  a  closely 
cropped,  rough,  much  trampled  piece  of  pasture.  Where  a 
year  ago  about  60  different  plants  had  grown  in  wild  exuber- 
ance, only  about  40  cropped  and  crippled  trampled  species  were 
to  be  found*  and  only  a  single  one  of  these  bloomed  and  flour- 
ished in  large  numbers,  the  formidable  armed  thistle,  Carduus 

*  About  30  species  of  the  list  of  August  16,  1899,  had  survived  fire  and  cattle 
and  about  10  species  were  found  that  do  not  appear  on  the  list  of  August  16, 
180.0. 


Langc :    REVEGETATIOX  OF  TRESTLE   ISLAND. 


629 


lanceolata.  It  has  occupied  nearly  one  third  of  the  section 
and  next  summer,  I  think,  the  thistle  will  keep  the  cows  off  the 
grass,  and  as  the  West  section  is  a  willow  island,  the  East  sec- 
tion will  be  a  thistle  island. 

During  September,  1900,  there  was  over  5  inches  of  rain  and 
now  water  once  more  surrounds  the  East  section  and  the  cows 
have  not  been  there  since  about  October  ist. 

The  West  Section  in  the  Season  of  1900. — The  lake  has  risen 
about  two  feet  since  September  i5th  this  year  and  the  south 
half  of  the  section  is  under  water.  Fire  and  cattle  have  thus 
far  never  invaded  this  section.  The  willows  are  thriving.  There 
are  probably  five  species  of  them,  but  Salix  amygdaloidcs 
forms  the  thicket.  There  are  a  few  thrifty  cottonwoods,  and 
aspens,  one  balsam  poplar,  and  one  slippery  elm,  Ulmus  fulva. 
Although  sumac,  hazel,  box-elder,  silver  maple,  wild  haw, 
flowering  dogwood  and  three  species  of  oak  grow  within  half 
a  mile  and  most  of  them  within  a  stone's  throw  of  the  island, 
not  one  individual  of  all  these  has  been  found  on  either  section 
of  Trestle  island. 

Last  spring  several  of  the  willow  species,  when  they  were 
only  about  24  months  old,  bloomed  'for  the  first  time  and  pro- 
duced fruit.  A  count  on  May  nth  revealed  38  staminate  indi- 
viduals and  27  pistillate  individuals  of  Salix  amygdaloides  in 
flower.  The  count  was  then  made  to  include  older  trees  of 
the  same  species  along  the  cycle  path  near  the  island  and  of  a 
total  of  135  trees  counted  74  bore  male  and  61  bore  female 
flowers.  Although  a  number  of  the  trees  were  measured  last 
year,  only  a  few  recent  measurements  of  October  12,  1900, 
can  be  given  here. 


Number. 

Diameter  at  4 
inches  above  the 
ground. 

Height 
measured  along 
the  stem. 

Growth  of  this 
season. 

i.  Salix  amgydaloides. 
2.  Salix  amygdaloides. 
3.  Salix  amygdaloides, 
4.  Populus  delloides. 
5.  Populus  deltoides. 
6.  Popultts  tremuloides. 
7.   Ulmus  fulva. 

3      inches 

In 

5fi 

i 
% 

15  ft.  2  in. 
16  ft.  IO  in. 
I8ft. 
14  ft.  3^  in. 
15  ft.  9  in. 
10  ft.  7  in. 
4ft.  Sin. 

7  ft.  2  in. 
Uncertain. 
7  ft.  3/2  in. 
6  ft.  9^  in. 
6  ft.  6  in. 
4  ft.  2  in. 
2  ft.  9  in. 

There  are  many  willows  on  this  section  of  the  size  of  those 
measured,  while  the  two  cottonwoods  and  the  aspen  measured 
are  the  largest  that  could  be  found  and  the  elm  is  the  only  indi- 
vidual present.  The  average  size  of  the  large  vigorous  Salix 


630  MINNESOTA    BOTANICAL    STUDIES. 

amy gdaloides  individuals  is  about  2^-3  inches  in  diameter  at 
four  inches  from  the  ground,  and  16  feet  in  height.  The  larg- 
est cottonwood,  No.  5,  shows  a  horizontal  spread  of  its  top  of 
2^  feet,  while  a  willow  (S.  amygdaloidcs}  near  it  spreads  its 
branches  8  feet  in  a  horizontal  diameter.  A  number  of  trees 
have  been  marked  and  their  study  as  well  as  that  of  the  whole 
island  is  being  continued.  The  willows  are  now  so  large  that 
for  some  time  a  flock  of  English  sparrows  regularly  roost  in 
them  and  they  seem  to  prefer  the  part  of  the  section  that  is 
flooded. 

ANIMAL  LIFE  OF  THE  ISLAND. 

Did  the  limits  of  this  paper  permit,  an  interesting  chapter 
might  be  added  under  this  heading.  Crayfish,  voles,  mice, 
and  muskrats  burrow  under  and  in  the  island,  since  the  sum- 
mer of  1898,  the  cottontails  resort  to  it,  and  in  August  of  1899 
some  minks  had  made  their  home  under  the  old  ties,  which 
once  formed  the  engineer's  pontoon.  Near  the  mink's  home  a 
song-sparrow  had  hatched  its  young  on  a  thistle  bush.  Some 
of  the  young  trees  on  the  West  section  have  been  infested  with 
the  spotted  willow  aphid,  Mclanoxanthus  saliciis,*  since  the 
summer  of  1899  and  their  secretions  attract  swarms  of  flies  and 
wasps.  That  the  frogs  are  there  is  self-evident,  but  I  also 
captured  a  fine  green  tree-frog,  and  a  bunch  of  prickly  cater- 
pillars of  the  morning-cloak  butterfly,  Euvanessa  antiopa,  found 
their  table  spread  on  the  willows. 


*  Identified  by  Dr.  Otto  Lugger. 


XXXVII.     VIOLET    RUSTS    OF    NORTH    AMERICA. 


J.  C.  ARTHUR  AND  E.  W.  D.  HOLWAY. 


A  rust  of  violets,  in  its  three  forms  of  aecidium,  uredo  and 
teleutospore,  is  common  throughout  North  America  upon  nearly 
all  indigenous  species  of  the  genus  Viola.  For  the  most  part 
it  belongs  to  a  single  species,  Puccinia  Violce  (ScnuM.)  DC., 
which  is  also  the  common  violet  rust  of  Europe  and  of  some 
other  regions.  This,  at  least,  is  the  conclusion  to  which  we 
have  arrived  after  a  rather  extended  study  of  considerable  ma- 
terial. Beside  the  one  common  rust  there  is  a  peculiar  sEcidium 
throughout  the  eastern  part  of  North  America,  and  one  species 
of  Puccinia  in  the  western  part,  both  distinctly  American. 

In  this  connection  we  desire  to  acknowledge  the  kindness  of 
the  New  York  Botanical  Garden,  the  Botanical  Department  of 
the  University  of  Illinois,  and  of  the  Iowa  State  College,  in 
loaning  material  from  their  herbaria,  and  to  extend  our  thanks 
to  their  representatives.  We  wish  also  to  thank  Dr.  J.  J.  Davis, 
of  Racine,  Wis.,  and  Mr.  E.  Bartholomew,  of  Rockport,  Kans., 
for  aiding  us  with  specimens  and  information.  We  are  further- 
more grateful  to  Mr.  Stewardson  Brown,  curator  of  the  herba- 
rium of  the  Philadelphia  Academy  of  Sciences,  for  the  privilege 
of  examining  material  in  the  Schweinitz  collection,  and  to  the 
custodians  of  the  Gray  Herbarium  of  Harvard  University  and 
of  the  New  England  Botanical  Club,  from  the  examination  of 
whose  phanerogamic  collections  of  Viola,  five  specimens  of  rust 
were  obtained  in  the  former  instance  and  three  in  the  latter. 

In  this  article  for  conciseness  we  have  used  in  addition  to 
the  usual  I,  II  and  III  for  designating  the  aecidium,  uredo  and 
teleutospore  stages,  the  sign  O  for  the  spermogonial  stage.  In 
citing  specimens  these  signs  are  put  into  large  type  when  the 
stage  is  present  in  abundance,  and  into  small  type  when  sub- 
ordinate and  in  small  amount. 

631 


632  MINNESOTA    BOTANICAL    STUDIES. 

^cidium  pedatatum  (ScH\v.)  nom.  nov. 
SYN.  : 

1834.    Cceoma     (sEcidium )     pedatatum     SCHW.      Trans. 

Amer.  Phil.  Soc.  4:   293,  no.  2885. 
1834.    Cceoma      (sEcidium)     sagittatum    SCHW.      Trans. 

Amer.  Phil.  Soc.  4:   293,  no.  2886. 
1672.  j&cidium  Maria-  Wilsoni    PK.   24th    Rep.    N.    Y. 

Mus.  for  1870  :  92. 

1874.  -dZcidium  Petersii^B.  &  C.   Grev.  3:   61. 
Exsicc.  : 

Carleton,  Ured.  Amer.  no.  28. 

Hetercecious,  inhabiting  species  of  Viola,  and  also  of  some 
other  genus  of  plants  not  yet  determined. 

0.  Spermogonia  preceding  the  ascidia. 

1.  ./Ecidia  hypophyllous,  seated  on  small,  pale,  circumscribed,  un- 
thickened  spots ;    cups  usually  sub-circinating,  small,  shallow  or  short 
cylindrical,  white,  border  narrow,  often  much  split  and  somewhat  re- 
curved;    spores  subglobose,  in  part  angular  from  compression,   epi- 
spore  thin,  minutely  verrucose,  n  to  18  //,  averaging  14, a. 

Throughout  the  United  States  east  of  the  Rocky  mountains, 
from  March  to  June  on  the  blades  of  various  species  of  violets, 
and  less  often  on  the  petioles,  pedicels  and  calyx.  Specimens 
have  been  examined  as  follows  : 

On  Viola  obliqua  HILL.  Pennsylvania  (Schweinitz),  Maine 
(Blake),  Indiana  (Arthur),  Illinois  (Seymour,  Arthur), 
Iowa  (Holway),  Kansas  (Bartholomew,  Carleton). 

On   Viola  ovata  NUTT.     New  Jersey  (Ellis). 

On  Viola  sagittata  L.  Pennsylvania  (Schweinitz),  Massa- 
chusetts (Asa  Gray). 

On  Viola  pedata  L.  Pennsylvania  (Schweinitz),  Alabama 
(Baker),  Iowa  (Holway). 

On  Viola  pedatifida  DON.  Kansas  (Kellerman),  Iowa 
Holway). 

On  Viola primulcef alia  L.  District  of  Columbia  (Greene), 
Mississippi  (Tracy). 

On   Viola  Nuttallii  PURSH.     Kansas  (Bartholomew). 

On  Viola  striata  AIT.  North  Carolina  (Biltmore  Herba- 
rium). 

On   Viola  tricolor  L.     Kansas  (Popenoe). 

The  form  here  described  is  undoubtedly  part  of  an  hetercecious 
species,  having  the  alternate  forms  possibly  upon  carices.  It 


Arthur  and  Hoi-way:    VIOLET  RUSTS  OF  NORTH  AMERICA.     633 

occurs  in  open  woodlands  from  the  last  of  March  to  the  begin- 
ning of  June,  but  is  rather  infrequent  and  local.  It  is  distin- 
gished  from  the  secidium  of  Puccinia  Violce  by  the  markedly 
smaller  cups  and  spores,  and  by  a  greater  tendency  to  form 
small,  round  groups  seated  on  the  mesophyll  of  the  blade, 
rather  than  on  the  midrib  or  veins.  The  peridium  is  usually 
conspicuously  white ;  the  cylindrical  elongation  noted  in  the 
description  by  Berkeley  and  Curtis  is  occasionally  to  be  seen, 
but  must  be  considered  as  accidental,  depending  upon  condi- 
tions of  growth.  The  same  kind  of  development  is  met  with 
in  other  forms  of  gecidia  (see  notes  on  J&cidium  -pulcherrimum 
Rav.  in  Bull.  Lab.  Nat.  Hist,  of  Univ.  Iowa  4:  399-400). 

Examination  of  specimens  in  the  Schweinitz  collection  at  the 
herbarium  of  the  Philadelphia  Academy  of  Sciences  shows  that 
both  species  which  he  described  as  new,  occurring  on  Viola 
•pcdata  and  V.  sagittata  respectively,  belong  here.  The  name 
coming  first  on  the  page  is  taken  as  the  authentic  name  of  the 
species.  The  Schweinitz  specimens  on  both  hosts  are  ample, 
but  neither  shows  the  Eecidia  in  so  characteristic  a  form  as  does 
the  specimen  in  the  same  collection  on  Viola  obliqua,  which 
Schweinitz  referred  to  sEcidium  Violarum.  The  last  has  very 
small,  compactly  clustered  aecidial  cups,  in  small  round  groups, 
while  both  the  former  have  the  cups  somewhat  larger,  and 
sparsely  scattered  or  solitary.  The  difference  in  habit  is  doubt- 
less due  to  the  influence  of  the  host  and  the  atmospheric  con- 
ditions affecting  the  development,  in  part,  combined  with  the 
accidental  distribution  of  the  infection  over  the  leaf  surface. 

Spermogonia  have  not  been  seen  by  the  writers,  but  Burrill 
(Par.  Fung.  111.,  p.  223)  states  that  they  precede  the  secidia,  al- 
though he  does  not  describe  them. 

Puccinia  Violas  (SCHUM.)  DC.     1815.     Flore  Francaise  6:62. 
SYN.  : 

1803.  jEcidium   Viola  SCHUM.  Fl.  Saell.  2:  224. 
1803.    Uredo   Viola  SCHUM.  Fl.  Saell.  2:  233. 
1805.  ^Ecidium   Violarum  DC.  Fl.  Franc.  2:  240. 
1825.  Puccinia   Violarum  LINK.  Sp.  Plant.  2:  80. 
1875.  Puccinia  hastatce  COOKE,  Grev.  3:  179. 
1888.    Puccinia    Fergussoni    hastatce   DsT.    Sacc.    Syll. 
Fung.  7:  682. 

1897.  Puccinia  densa  D.  &  H.   Hedw.  26:  298. 

1898.  Dicceoma  Violce  KUNTZE,  Rev.  Gen.  PL  3:  471. 


634  MINNESOTA    BOTANICAL    STUDIES. 

Exsicc.  : 

Jaczewski,   Komarov,  Tranzschel,  Fungi  Ross.   no.  i6m. 

Ellis   and  Everhart,  N.   Am.   Fungi,   nos.    254 In,    1007 T, 
241 1  m. 

Seymour  and  Earle,  Econ.  Fungi,  no.  456  m. 

Thueman,  Myc.  Univ.,  no.  430 J. 

Linhart,  Fungi  Hung.,  no.  33  T. 

Krieger,  Fungi  Sax.,  nos.  no1,  in  HI,  472  n,  561 1,  562  u, 
1006 1. 

Rabenhorst,  Fungi  Europ.,  no.  2I941. 

Sydow,  Mycotheca  Marchica,  no.  468 m. 

Sydow,  Ured.  Exsicc.,  nos.  33  m,  82  I,  117",  286",  335  x, 
378",  640 J,  935  l,  1136  m,  1184 J,  1229 J. 

Shear,  N.  Y.  Fungi,  no.  321  m. 

Vize,  Fungi  Brit.,  no.  77 x,  ii2m. 
Autaecious,  inhabiting  species  of    Viola. 

0.  Spermogonia  preceding  or  accompanying  the  aecidia,    amphi- 
genous,  punctate,  honey  yellow ;  spores  elliptical  or  nearly  globose,  3 
to  5  fi  broad  by  4  to  6.5  /j.  long. 

1.  yEcidia  hypophyllous  and  also  on  petioles,  pedicels  and  calyx, 
substratum  moderately  thickened,  in  indefinite  and  irregular  clusters, 
often  covering  nearly  the  whole  leaf,  especially  noticeable  on  the  veins 
and  stalks,  crowded ;  cups  broad  and  low,  rather  coarsely  lacerate  and 
irregularly  recurved;  spores  subglobose,  somewhat  angular  from  com- 
pression, minutely  verrucose,  14  to  i8//  broad  by  15  to  22  ;j.  long  (Eu- 
ropean) or  16  to  2O/J.  broad  by  18  to  26 /JL  long  (American)  ;  wall  thin. 

II.  Uredo  chiefly  hypophyllous;  sori  at  first  in  small  groups  on  dis- 
colored spots,  later  sparsely  and    indefinitely  scattered,   soon  naked, 
cinnamon  brown,  pulverulent ;   spores  subglobose,  echinulate,    17  to 
28^1  in  diameter,  brownish  yellow;  wall  varying  from  thin  to  thick; 
pores  four,  equatorial. 

III.  Teleutosori  hypophyllous,  indefinitely  scattered,  round,  small, 
soon  naked,    pulverulent,    chocolate   brown ;    spores    brown,  usually 
broadly  elliptical,  less  often  oblong-ovoid  or  irregular,  slightly  or  not 
at  all  constricted  at  the  septum,  smooth,  or  finely  tuberculate  especially 
on  the  upper  half,  15  to  23, a  broad  by  21  to  30  ,u  long  (European),  or 
16  to  26 11.  broad  by  28  to  44/>-  long  (American)  ;  apex  obtuse,  some- 
what thickened,  with  a  pale  and  broad  apicuous,  while  a  similar  pro- 
jection often  occurs  on  one  side  of  the  lower  cell   near  the  septum ; 
pedicel  hyaline,  fragile,  not  as  long  as  the  spore,  somewhat  deciduous. 

Throughout  North  America,  occurring  upon  nearly  every  indi- 
genous species  of  Viola.    Spermogonia  and  ascidia  from  April  to 


Arthur  and  Holivay:    VIOLET  RUSTS  OF  NORTH  AMERICA.     635 

June,  uredo  and  teleuto  stage  from  June  to  November  and  later 
along  the  Gulf  region.  Specimens  of  American  origin  have 
been  examined  as  follows  : 

On   Viola  obliqua  HILL.      1879?  Newton,  Mass.  III.  (  W.  G. 

Farlow);   1882,  Decorah,  Iowa  III  (E.   W.  D.  Holway); 

1883,  Madison,  Wis.  I  (L.  H.  Pammel);  1885,  Oregon, 

111.  Ill  (M.  B.  Watte);   1885,  Decorah,  Iowa,  ii.  Ill  (E.  W. 

D.Hohvay);  1886,  Manhattan,  Kans.  I  and  ii.  Ill  ( W. 

A.    Kellerman);     1890,    Greencastle,    Ind.    O.   I   (J.    C. 

Arthur};   1893,   Terre   Haute,   Ind.    I  (J.    C.    Arthur); 

1897,  Auburn,  Ala.  I  (Ala.  BioL  Surv.);   1898,  Lafay- 
ette, Ind.  O.  I  (/.  C.  Arthur). 
On  Viola  rotundifolia  MICHX.     1890,  Bradley,  Me.  II  (F.  L. 

Harvey};    1891,  Gunflint  lake,  Minn.    i.    II.   Ill  (L.  S. 

Cheney);    1899,    St.    Louis   river,  Wis.  i.   II.   iii    (L.    S. 

Cheney. 
On   Viola  renifolia  A.  GRAY.     1896,  Orono,  Me.  I   (E.   D. 

Merrill). 
On  Viola  septentrionalis  GREENE.     1898,  Ottawa,  Ont.  I  (J. 

M.  Maconn). 
On  Viola   blanda  WILLD.     1885,   Lansing,  Mich.  ii.  Ill  (J. 

C.  Arthur);  1886,  Vermilion  lake,  Minn.  II.  iii  (E.   W. 

D.  Holivay');    1892,    Oakland  Co.,  Mich.  II.  iii  (G.  H. 
Hicks) ;  1899,  Isle  au  Haut,  Me.  ii.  Ill  (J.  C.  Arthur). 

On  Viola  primulcefolia  L.  1892,  Lake  City,  Fla.  II.  iii  (P. 
H.  Rolfs);  1899,  Isle  au  Haut,  Me.  ii.  Ill  (/.  C.  Arthur]. 

On  Viola  lanceolata  L.  1897,  Sault  Ste.  Marie,  Mich.  II. 
(E.  T.  Harper);  1899,  Isle  au  Haut,  Me.  ii.  Ill  (/.  C. 
Arthur). 

On  Viola  cognata  GREENE.  1894,  North  Yakima,  Wash.  ii. 
Ill  (C.  V.  Pi^er};  1894,  Sisson,  Calif,  ii.  Ill  (  W.  C.  Bias- 
dale)  . 

On  Viola  adunca  SMITH.  1887,  Provo,  Utah,  ii.  Ill  (S.  M. 
Tracy) ;  1889,  Deer  Lodge,  Mont.  I  (F.  D.  Kelsey);  1892, 
Pullman,  Wash.  I  and  ii.  Ill  (W.  7?.  Hull);  1894,  Pull- 
man, Wash.  II.  iii  ( C.  V.  Piper);  1894,  Pine  Creek, 
Calif,  i.  II.  iii  (F.  P.  Nutting);  1894,  Sisson,  Calif.  I  (E. 
W.  D.  Holway);  1894,  Sisson,  Calif.  II.  iii  (W.  C.  Bias- 
dale);  1899,  Gunnison  Co.,  Colo.  ii.  Ill  (E.  Bartholomew). 

On  Viola  glabella  NUTT.  1885,  Falcon  Valley,  Wash.  ii. 
Ill  (W.  JV,  Suksdorf);-Lfy$,  Skamania  Co.,  Wash.  Ill 


636  MINNESOTA    BOTANICAL    STUDIES. 

(W.  N.  Suksdorf};    1897,  Bingen,  Wash.   I.  II.  Ill  (W. 

JV.  Suksdorf}. 
On   Viola  ocettata  TORR.  &  GR.      1894,  Ukiah,  Calif,  ii.  Ill 

(E.   W.  D.  Hoi-way). 
On   Viola  Montanensis  RYDB.      1888,  Helena,  Mont.  II.   Ill 

(F.  D.   Kelsey};    1897,   Spanish   Basin,  Mont.  ii.  Ill    (P. 

A.  Rydberg);   1898,  Montesanes,   Wash.    ii.    Ill  (A.    A. 
Heller). 

On   Viola  Canadensis  L.      1889,  Bozeman,  Mont.  O.  I  (Mrs. 

Alder  son}. 
On    Viola  striata    AIT.      1882,  Pine   Hills,  111.  I.  ii.  Ill  (A. 

B.  Seymour)',    1882,  Carbondale,    111.  I  (A.  B.  Seymour); 
1893,  Ell  River  Falls,  Ind.  ii.  Ill  (L.  M.  Underwood). 

On  Viola  rostrata  PURSH.  1886,  Syracuse,  N.  Y.  I  (L. 
M.  Under-wood). 

On  Viola  Labradorica  SCHRANK.  1883,  Granville,  Mass. 
ii.  Ill  (A.  B.  Seymour};  1885,  Racine,  Wis.  ii.  Ill  (/.  /. 
Davis);  1886,  Racine,  Wis.  I  (J.  J.  Davis);  1888,  Racine, 
Wis.  ii.  Ill  (J.  J.  Davis);  1896,  St.  Remi,  Quebec,  ii. 
Ill  (  Wm.  Stuart}. 

On  Viola  arenaria  DC.  1880,  Franklin  Falls,  N.  H.  II 
(Mrs.  Harrison). 

On  Viola  hastata  L.  1820?  Salem,  N.  C.  I  (L.  v.  Schivei- 
nitz}. 

On  Viola  -pubescens  AIT.  1882,  Decorah,  Iowa,  I  (E.  W. 
D.  Hoi-way);  1883,  Madison,  Wis.  ii.  Ill  (L.  H.  Pammel); 
1883,  Devil's  Lake,  Wis.  Ill  (L.  H.  Pammel);  1883, 
Decorah,  Iowa  ii.  Ill  (E.  W.  D.  Holway};  1885,  Madi- 
son, Wis.  ii.  Ill  (A.  B.  Seymour);  1885,  Racine,  Wis.  ii. 
Ill  (/.  /.  Davis};  1886,  Racine,  Wis.  O.  I  (/.  /.  Davis}; 
1886,  Decorah,  Iowa,  O.  I  (E.  W.  D.  Holway);  1889, 
Greensburg,  Ind.  I  (J.  C.  Arthur);  1898,  Lafayette,  Ind. 
O.  I  (/.  C.  Arthur);  1898,  Pownal,  Vt.  O.  I  (/.  R. 
Churchill). 

On  Viola  scabriuscula  (T.  &  G.)  SCHW.  1898,  Franconia, 
N.  H.  I  (E.  &  C.  E.  Faxon};  1898,  Masardis,  Me.  I  (M. 
L.  Fernald). 

On  Viola  sp.  1884,  Jamestown,  N.  D.  II.  Ill  (A.  B.  Sey- 
mour); 1893,  Three  Lakes,  Wis.  I  (J.  J.  Davis};  1895, 
Mt.  Washington,  N.  H.  ii.  Ill  (E.  T.  Harper);  1897, 
SaultSte.  Marie,  Mich.  I  (E.  T.  Harper};  1899,  Pachuca, 
Mexico  II.  Ill  (E.  W.  D.  Hoi-way}. 


Arthur  and  Hotway :   VIOLET  RUSTS  OF  NORTH  AMERICA.     637 

This  species  shows  remarkable  variability,  especially  in  size 
of  the  spores,  and  in  the  thickness  and  markings  of  their  walls. 
These  differences  come  out  most  strikingly  when  comparing  a 
series  of  specimens.  There  is  also  considerable  variability  in 
the  form  of  the  spores,  as  shown  in  Figs.  5  and  6,  both  taken 
from  the  same  specimen,  but  it  is  such  as  one  may  expect  to 
find  in  many  other  species  of  rusts. 

If  almost  any  specimen  of  violet  rust  be  compared  critically 
with  an  European  specimen,  the  greater  size  of  the  teleutospores 
in  the  former  is  likely  to  attract  attention.  This  is  the  basis  of 
the  species  proposed  by  Cooke,  Puccinia  hastatce.  A  specimen 
collected  by  W.  C.  Blasdale  on  Viola  cognata  at  Sisson,  Calif. 
(Figs.  5  and  6),  has  been  compared  with  the  type  material  of 
P.  hastatce  at  the  Kew  Herbarium,  through  the  kindness  of  Mr. 
Massee,  and  found  to  agree  very  closely.  The  greater  size  of 
the  teleutospore  in  the  American  material  generally  is  notice- 
able, and  this  difference  extends  to  the  uredo  and  ascidial  stages 
as  well,  although  not  usually  so  pronounced.  In  the  case  of 
an  ascidium  on  Viola  pubesccns,  collected  at  Decorah,  Iowa,  in 
1882,  Farlow  has  made  the  comment: — "Spores  somewhat 
larger  than  in  the  European  specimens ;  this  may  be  the 
^•Ecidium  of  Puccinia  hastata  Cke."  (on  the  label  in  Ellis'  N. 
Amer.  Fungi,  no.  1007).  If  size  of  spores  is  to  be  taken  as 
valid  basis  for  separating  species,  there  is  no  question  that  the 
American  form  shows  a  strong  claim  to  rank  as  autonomous. 
The  claim,  even  on  that  assumption,  is  much  weakened,  how- 
ever, by  the  great  range  between  the  smallest  and  largest  of 
the  American  specimens,  indicating  a  decided  capacity  for  vari- 
ability rather  than  a  fixed  form. 

A  peculiarity  of  the  American  violet  rust,  that  in  the  case  of 
European  specimens  we  have  not  seen  mentioned  and  have  not 
observed,  is  the  frequently  tuberculate  sculpturing  of  the  teleu- 
tospores. Burrill  (Paras,  fungi  111.  :  174)  makes  this  a  diag- 
nostic character,  but  in  a  wide  series  of  specimens  one  does  not 
always  meet  with  it.  With  most  observers  the  spores  would 
generally  be  rated  as  smooth  like  the  European  form.  The 
true  character  of  surface  markings  can  be  best  studied  by  ob- 
serving the  spores  without  addition  of  fluids.  In  this  way  it  is 
easy  to  see  that  the  markings  are  small  papillae,  sparingly  dis- 
tributed, and  chiefly  appearing  on  the  upper  half  of  the  spore. 
A  closer  study  reveals  the  interesting  fact  that  when  no  eleva- 


638  MINNESOTA    BOTANICAL    STUDIES. 

tions  above  the  general  surface  of  the  spore  can  be  detected, 
there  may  yet  be  observed  almost  the  usual  appearance  when 
the  spore  is  examined  in  face  view.  Instead  of  papillae,  their 
places  seem  to  be  supplied  by  translucent  dots.  Now  the  most 
interesting  outcome  of  this  study  is  the  observation  that  the 
European  specimens,  while  having  what  are  always  rated  as 
smooth  teleutospores,  yet  show  when  looked  at  dry  and  in  face 
view,  the  same  appearance  of  translucent  dots  and  in  the  same 
abundance  and  distribution  as  do  American  specimens.  The 
American  form,  therefore,  simply  accentuates  characters  that 
are  primitive  in  the  trans-Atlantic  form. 

The  uredospores  also  have  interesting,  but  less  significant, 
characters  showing  variability.  As  a  rule  they  do  not  much 
exceed  in  size  those  from  European  specimens,  although  the 
tendency  toward  largeness  is  apparent.  But  in  many  American 
specimens  the  walls  are  greatly  thickened  (compare  uredospores 
in  Figs.  5  and  7),  and  give  a  striking  appearance  under  the 
microscope.  These  thick-walled  uredospores  are  sometimes 
small,  and  sometimes  large.  They  occur  on  various  species  of 
violets  and  range  from  the  Atlantic  coast  to  the  Pacific.  Fine 
illustrations  are  found  in  material  on  Viola  -primulcBfolia  col- 
lected at  Isle  au  Haut,  Me.  (Fig.  8),  the  uredospores  being 
small,  and  on  Viola  glabella  collected  at  Bingen,  Wash.  (Fig. 
7),  the  uredospores  being  large.  After  all,  the  form  is  only  oc- 
casionally met  with.  If  it  is  an  adaptation  to  some  particular 
environment,  it  is  difficult  to  see  what  that  may  be.  The  form 
on  Viola  -primula/oHa  from  Maine  was  collected  within  a  few 
hundred  feet  of  the  open  ocean  among  rocks,  yet  in  the  same 
situation  and  intermixed  grew  Viola  lanceolata  with  rust  show- 
ing uredospores  having  almost  normally  thin  walls. 

It  is  possible  that  these  interesting  variations  belong  in  some 
way  to  obscure  species,  but  our  study  has  shown  no  morpho- 
logical boundaries.  It  is  more  likely  that  they  indicate  races, 
or  possibly  so-called  biological  species.  A  well-directed  series 
of  cultures  would  undoubtedly  yield  important  results. 

A  few  words  regarding  the  American  synonymy  may  be 
helpful.  Cooke  described  Puccinia  hastattz  in  the  third  volume 
of  Grevillea  from  material  collected  in  Maine  by  E.  C.  Bolles. 
The  host  was  Viola  hastata.  The  uredo  and  teleuto  stages  are 
described,  but  the  only  distinctive  characters  are  the  measure- 
ments which  are  given  as  20-22  //  for  the  uredospores,  and 


Arthur  and  Holway :    VIOLET  RUSTS  OF  NORTH  AMERICA.     639 

20-25  p  by  35-40;*  for  the  teleutospores.  The  name  has  been 
very  little  used  by  American  or  other  botanists.  The  assign- 
ment of  the  name  to  a  place  under  the  wholly  distinct  Puccinia 
Fergussoni,  as  done  by  De  Toni  in  the  seventh  volume  of 
Saccardo's  Sylloge  fungorum,  was  far  from  being  a  shrewd 
guess  as  to  its  relationship.  It  is  even  more  inexplicable  how 
Dietel  could  have  fancied  a  resemblance  to  Puccinia  Fergussoni 
in  the  type  material  of  his  Puccinia  dcnsa.  His  material  of  the 
latter  species  was  collected  in  1895  by  W.  N.  Suksdorf  in  the 
State  of  Washington.  It  was  on  Viola  glabella,  and  yielded 
only  teleutospores.  The  characters  which  he  drew  up  for  the 
proposed  new  species  agree  perfectly  with  those  of  large-spored 
forms  of  Puccinia  Viola.  Beside  the  type  collection  we  have 
examined  material  on  the  same  host  from  other  localities  in  the 
same  State,  and  secured  by  the  same  collector.  This  ample 
material  includes  all  three  spore  stages,  and  leaves  no  doubt  of 
the  identity  of  P.  densa  Diet,  with  P.  Viola. 

All  the  hosts  of  the  specimens  cited  under  Viola  Montanensis, 
and  part  of  those  under  V.  adunca,  have  been  determined  or 
verified  by  Dr.  P.  A.  Rydberg  of  the  New  York  Botanical  Garden. 

An  error  in  the  thirteenth  volume  of  Saccardo's  Sylloge  fun- 
gorum,  page  1313,  should  be  pointed  out  here.  Puccinia 
Maria-  Wilsoni  Clint,  is  said  to  occur  on  Viola  cucullata  Ait. 
and  V.  delphinifolia  Nutt.  The  error  is  due  to  a  confusion  of 
names.  ^Ecidium  Maria-  Wilsoni  Peck  is  found  on  these 
hosts  but  Puccinia  Maria-  Wilsoni  Clinton  is  only  found  on 
Claytonia,  and  both  species  are  widely  different  from  Puccinia 
Viola. 

Puccinia  effusa  D.  &  H.     1895.     Erythea  3:  81. 

I.  yEcidia  amphigenous,  but  chiefly  hypophyllous,  in  large  indefi- 
nite clusters,  often  covering  much  of  the  leaf,  noticeably  extending 
along  the  veins  and  petioles ;  substratum  somewhat  thickened ;  cups 
broad  and  low,  border  white,  irregularly  and  coarsely  lacerate,  some- 
what recurved  ;  spores  subglobose,  somewhat  angular  from  compres- 
sion, minutely  verrucose,  20—27  ;j.  in  diameter. 

III.  Teleutosori  for  the  most  part  arising  from  the  cups  of  the 
aecidia,  uncovered,  elliptical,  nearly  black ;  spores  dark  brown,  ellip- 
tical or  oblong,  slightly  if  at  all  constricted  at  the  septum,  inconspicu- 
ously verrucose,  23-317^  broad  by  37-50/4  long;  wall  moderately 
thick ;  apex  rounded,  usually  not  thickened ;  base  rounded  or  occa- 
sionally slightly  narrowed;  pedicel  hyaline,  deciduous. 


640  MINNESOTA    BOTANICAL    STUDIES. 

The  western  coast  of  the  United  States  in  spring  and  early 
summer.  The  following  specimens  have  been  examined,  the 
first  being  the  type  collection  : 

On  Viola  lobata  BENTH.  1894,  Dunsmuir,  Calif.  I.  iii  (E. 
W.  D.  Holway). 

On  Viola  Nuttallii  PURSH.  1897,  Falcon  Valley,  Wash.  I. 
iii  (  W.  N.  Suksdorf}. 

The  species  is  very  characteristic.  It  probably  possesses 
sperrnogonia,  but  they  have  not  yet  been  observed ;  it  is,  how- 
ever, without  uredo,  although  erroneously  included  in  the  orig- 
inal description  of  the  species.  Two  specimens  are  cited 
(Erythea  3 :  82)  as  type  material,  the  first  on  Viola  lobata,  re- 
ferred to  above,  and  the  second  on  Viola  ocellata.  The  latter 
specimen  has  been  examined  and  proves  to  be  Puccinia  Violce, 
and  is  cited  above  under  that  species.  The  original  description 
of  the  species  is  accordingly  emended  to  omit  the  supposed 
uredo. 

The  difference  in  the  shape  of  the  spores,  shown  in  the  pho- 
tographs of  spores  taken  from  the  two  hosts,  is  doubtless  due 
to  some  accident  of  growth,  such  as  more  or  less  compression 
in  the  young  sorus,  and  is  without  diagnostic  value.  It  is  of 
the  same  nature  as  the  difference  shown  in  two  mountings  from 
the  same  collection  of  P.  Viola  on  V.  cognata.  The  more 
regular  spores  are  to  be  accepted  in  each  case  as  the  normal 
development  under  favorable  conditions. 

Four  additional  species  of  violet  rusts  occur  in  Europe,  and 
it  is  possible  that  they  may  eventually  be  found  in  this  country. 
Uredo  alpestris  Schroet.  inhabits  Viola  biflora  L.,  and  as  this 
host  is  a  native  of  the  Rocky  Mountains,  the  rust  may  possibly 
accompany  it.  Puccinia  alpina  Fckl.  also  occurs  on  V.  bijlora 
L.,  and  P.  Fergussoni'Qvfa.  &  Br.  occurs  on  V.  -palustris  L., 
and  V.  mirabilis  L.,  and  V*  epipsila  Led.  Both  of  these 
species  belong  to  the  section  of  Micropuccinia,  and  are  notably 
distinct  from  the  other  rusts  on  violets.  A  specimen  of  rust 
collected  by  Marcus  E.  Jones  at  San  Diego,  Calif.,  in  1882, 
was  erroneously  referred  to  the  latter  species,  and  distributed 
by  him  as  No.  3040.  Puccinia  cegra  Grove  is  an  autoecious 
species  found  on  the  cultivated  pansy  and  its  close  relatives. 
It  will  probably  be  brought  to  America  after  a  time  through 
commercial  channels,  as  the  rusts  of  asparagus,  chrysanthe- 
mum, carnation,  hollyhock,  and  of  some  other  cultivated  plants 


Arthur  and  Holway :    VIOLET  RUSTS  OF  NORTH  AMERICA.     641 

have  been.  As  the  hosts  of  these  four  species  of  violet  rusts 
are  found  in  this  country,  the  rusts  may  not  unreasonably  be 
expected  also. 

EXPLANATION  OF  PLATE  XLVII. 

All  figures  photographed  and  engraved  to  the  same  scale,  x  250 
diameter.  Each  millimeter  on  the  plate  equals  4/z  of  original  dimen- 
sions. 

Figures  i,  2  and  3.  Puccinia  Viola  showing  three  sizes  of  teleuto- 
spores :  i,  on  V.  elatior  from  Germany,  Sydow's  Uredineen,  No.  33  ; 
2,  on  V.  adunca  from  Gunnison,  Colo. ;  3,  Viola  sp.  from  Mexico, 
Holway  No.  3573. 

Figures  4,  5  and  6.  Puccinia  Violce  showing  especially  large  teleu- 
tospores :  4,  on  V.  blanda  from  Lansing,  Mich. ;  5  and  6,  both  on  V. 
cognata  from  Sisson,  Calif.,  taken  from  different  sori. 

Figures  7  and  §•  Puccinia  Violce  showing  especially  thick-walled 
uredospores  :  7?  on  ^  glabetta  from  Bingen,  Wash. ;  8,  on  V.  primu- 
Icefolia  from  Isle  au  Haut,  Me. 

Figures  9  and  10.  Puccinia  effusa  showing  regular  and  irregular 
spores ;  9,  on  V.  Ntdtattii  from  Falcon  Valley,  Wash. ;  10,  on  V. 
lobata  from  Dunsmuir,  Calif. 


VOL.11.  MINNESOTA  BOTANICAL  STUDIES.  PART 


V. 


PLATE  XLVII. 


THE    MELWTVPf    PRINTINO    CO.,    BOSTON. 


XXXVIII.  OBSERVATIONS   ON   THE  EMBRYOGENY 
OF    NELUMBO. 


H.   L.   LYON. 


INTRODUCTION. 

The  peculiar  and  seemingly  inconsistent  characters  of  JVe- 
himbo  have  given  rise  to  a  variety  of  opinions  regarding  its 
proper  systematic  position  and  in  attempts  to  settle  the  points  in 
dispute  the  plant  has  again  and  again  been  subjected  to  careful 
investigation,  the  recorded  observations  forming  a  considerable 
literature. 

In  anatomy  the  plant  seems  to  conform  more  nearly  to  the 
type  of  the  Monocotyledons,  as  in  fact  do  all  the  Nymphaeaceae, 
the  vascular  bundles  being  closed  and  irregularly  placed  through 
the  stem.  On  the  other  hand  the  large  peltate  leaves  with  their 
reticulate  venation  are  perhaps  more  suggestive  of  a  dicoty- 
ledonous plant,  while  the  flowers  might  easily  belong  to  one  of 
either  class.  Thus  far  can  investigators  agree,  but  the  anoma- 
lous character  of  the  fruit  has  made  it  a  subject  for  controversy, 
and  the  interpretations  offered  are  numerous  and  at  great  vari- 
ance. Briefly  stated  the  fruit  presents  the  following  peculiari- 
ties. Each  carpel  of  the  apocarpous  gynoecium  contains  a 
single  ovule  and  matures  as  a  spherical  one-seeded  fruit.  The 
thick  sclerenchymatous  pericarp  lined  by  the  thin  testa  is  closely 
filled  by  two  large  white  fleshy  bodies  hemispherical  in  shape 
and  joined  to  each  other  at  the  stigmatic  end  of  the  pericarp. 
In  an  elongated  oval  chamber  formed  by  opposed  concavities 
in  the  inner  surfaces  of  these  fleshy  bodies,  and  attached  to 
them  at  their  point  of  continuity  is  a  green  structure — a  stem 
bearing  a  large  and  small  leaf  and  an  apical  bud  containing 
two  more.  The  free  leaves  are  fully  formed  and  already  green 
and  together  with  the  stem  are  enclosed  by  a  thin  delicate 
structureless  membrane.  Imbedded  in  the  common  tissue  of 
the  two  fleshy  bodies  opposite  the  insertion  of  the  stem  of  the 

643 


644  MINNESOTA    BOTANICAL    STUDIES. 

green  structure  is  found  a  vestigial  radicle.  Upon  the  germi- 
nation of  the  seed  the  fleshy  bodies  remain  within  the  pericarp 
wall  while  the  green  structure  develops  into  the  extraseminal 
plant  body.  The  radicle  does  not  function,  the  first  roots 
springing  from  the  stem  of  the  green  structure. 

The  first  careful  description  of  the  seed  seems  to  have  been 
given  by  Gaertner  (1788,  73-74).  He  termed  the  large  fleshy 
bodies  the  vitellus  which  he  considered  a  transition  between  the 
endosperm  and  cotyledon.  The  green  structure  he  considered 
the  embryo  and  in  Nymphcea  he  describes  it  as  monocotyledon- 
ous,  but  of  Nelumbo  he  says  :  "  Ambigit  Nelumbo  inter  plantas 
mono-&  dicotyledones :  nom  ad  posteriores,  ex  fabrica  seminis, 
omnino  spectare  videtur  ;  sed  verissime  ad  priores  pertinet,  quum 
constantissime  unicum  duntaxat  sub  germinatione  promat  foli- 
olum,  nee  alterum  prodeat,  donee  prius  penitus  evolutum  & 
super  aqua  explicatum  sit :"  Jussieu  (1789,  68,  453)  considers  the 
green  structure  a  monocotyledonous  embryo  and  describes  the 
large  fleshy  bodies  as  endosperm.  Poiteau  ('09,  382,  383)  in- 
terprets the  large  fleshy  bodies  as  cotyledons  and  the  membrane 
as  a  stipule,  but  denies  the  presence  of  a  radicle.  Mirbel  ('09) 
accepts  Poiteau's  interpretation  of  the  large  fleshy  bodies  but 
notes  the  presence  of  a  radicle.  Richard  ('n)  describes  the 
embryo  as  monocotyledonous.  The  little  sac  which  surrounds 
the  green  structure  he  considers  a  reduced  cotyledon  and  the 
large  fleshy  bodies  an  outgrowth  of  the  radicle.  Mirbel  ('i5» 
59,  60, footnote]  writes:  "Je  ne  suis  pas  eloigne  de  croire  que 

le  Piper,  le  Saurus ,  le  Nymphgea,  le  Nelumbo  et  peutetre 

quelques  autres  genres  que  Ton  regarde  mal-a-propos  comme 
Monocotyledons,  doivent  prendre  place  non  loin  les  uns  des 
autres,  parmi  les  Dicotyledons,  dans  la  serie  des  families  natur- 
elles."  MirbePs  declaration  seems  to  have  settled  the  question 
as  to  the  character  of  the  large  fleshy  bodies,  his  interpretation 
having  been  generally  accepted  except  by  Barthelemy  ('76)  who 
asserts  :  (i)  that  the  green  structure  is  the  one  which  arises  in  the 
embryo-sac  and  hence  is  to  be  considered  as  the  embryo,  (2) 
that  the  two  fleshy  bodies  imitating  cotyledons  arise  through  the 
division  of  the  exosperm. 

Concerning  the  little  colorless  sac  which  surrounds  the  green 
structure,  however,  no  opinion  seems  to  have  been  given  which 
could  meet  with  general  approval.  In  addition  to  those  above 
cited  De  Candolle  (J2i)  considered  it  a  stipule,  Brongniart  ('27) 


Lyon  :  OBSERVATIONS  ON  EMBRYOGENY  OF  XELUMBO.        645 

the  embryo-sac,  Trecul  ('54,  2)  a  concretion  of  a  homogeneous 
substance  containing  numerous  small,  acicular  crystals,  and 
finally  Wigand-Dennert  ('87)  in  a  monograph  on  Nehunbnini 
speciosum  wrote  :  "  Es  ist  unzweifelhaft  der  Ueberrest  von  Endo- 
spermzellen,  welche  sich  in  der  Hohlung  zwischen  den  Cotyle- 
donen  erhalten  und  gleichsam  auf  der  Oberflache  der  Plumula 
niedergeschlagen  und  dadurch  das  Ansehen  einer  Membran 
erhalten  haben." 

From  the  above  survey  of  literature  it  will  be  seen  that  mod- 
ern knowledge  of  the  seed  is  based  on  investigations  made  upon 
mature  or  nearly  mature  fruits.  That  alone  which  will  deter- 
mine the  correct  interpretation  of  the  structures  (Y.  e.,  a  knowl- 
edge of  their  origin)  is  entirely  wanting.  It  was  in  recognition 
of  this  fact  that  the  present  embryological  study  was  undertaken. 
In  a  preliminary  note,  published  in  Science,  the  more  important 
conclusions  were  announced,  i.  The  embryo  of  Nelumbo  is 
genuinely  monocotyledonous  in  its  development.  The  plumule 
arises  laterally  and  at  first  there  is  but  one  cotyledon  which  later 
bifurcates  to  form  the  two  fleshy  bodies.  2.  The  membrane 
surrounding  the  plumule  is,  as  conjectured  by  Wigand,  a  true 
endosperm  arising  within  the  embryo-sac. 

COLLECTION  AND  METHODS. 

The  material  for  the  investigation  in  hand  was  collected  by 
the  author  in  August,  1899,  and  August,  1900,  in  southeastern 
Minnesota  while  working  on  the  botanical  survey  of  that  region. 
There,  Nelumbo  lutca  grows  in  great  luxuriance  in  the  sloughs 
and  bayous  of  the  Mississippi  river,  the  soft  muddy  bottoms 
and  quiet  waters  of  these  bayous  affording  an  ideal  habitat  for 
the  plant,  where  it  often  forms  beds  many  acres  in  extent  to  the 
exclusion  of  other  vegetation. 

In  collecting,  each  carpel  was  removed  from  the  torus,  the 
lower  end  cut  away  and  the  upper  portion  with  the  attached 
ovule  placed  in  the  fixing  fluid.  One-half  and  one  per  cent, 
chromic  acid  and  chrom-acetic  acid  were  used  as  killing  re- 
agents. The  material  was  thoroughly  washed  and  passed  gradu- 
ally into  70  per  cent,  alcohol  in  which  condition  it  was  brought 
into  the  laboratory.  The  ovules  were  imbedded  in  paraffine  and 
serial  sections  cut  with  a  Minot  microtome.  The  sections  were 
stained  on  the  slide,  a  variety  of  stains  being  used ;  the  photo- 
micrographs and  drawings,  however,  which  accompany  the 


646  MINNESOTA    BOTANICAL    STUDIES. 

present  paper  were  made  from  sections  stained  with   aniline- 
water-safranin  or  acid  fuchsin. 

At  the  present  time  the  material  at  hand  is  insufficient  for  the 
full  demonstration  of  certain  points,  especially  in  the  develop- 
ment of  the  embryo-sac.  Therefore,  the  present  paper  will  be 
limited  to  a  discussion  of  the  embryogeny,  the  development  of 
the  embryo-sac  and  the  cytology  of  reproduction  being  reserved 
for  future  treatment. 

RESEARCH. 

The  embryo-sac  is  ovoid  at  its  micropylar  end  and  tapers  down 
to  a  narrow  attenuation  which  extends  deeply  into  the  nucellus. 
As  the  embryo-sac  matures  the  nucellar  tissue  directly  in  con- 
tact with  it  begins  to  break  down.  There  also  appears  in 
the  antipodal  region  a  club-shaped  cellular  structure  which 
obliterates  the  lower  portion  of  the  sac  (Figs,  i  and  2).  The 
precise  origin  of  this  peculiar  antipodal  body  remains  yet  to  be 
determined.  To  facilitate  description  the  embryogeny  may  be 
considered  in  a  series  of  stages. 

Stage  A.  Spherical  stage. — The  young  embryo  is  usually 
found  in  the  upper  end  of  the  embryo-sac  among  the  debris  re- 
sulting from  the  disintegration  of  the  synergidas.  The  sur- 
rounding protoplasm  makes  it  difficult  to  observe  the  first 
division  of  the  oosperm  and  at  the  present  time  the  youngest 
embryo  which  can  be  described  with  certainty  is  one  in  the 
eight-celled  stage  represented  in  Fig.  3.  As  seen  here  the  embryo 
is  nearly  spherical,  no  suspensor  being  evident,  although  there 
may  be  one  present  but  obscured  by  the  disorganizing  synergid 
"  a."  This  is  not  probable,  however,  judging  from  the  position 
of  the  first  wall  in  the  segmenting  embryo.  If  a  suspensor  cell 
is  cut  off  at  all,  it  very  early  loses  its  identity  as  there  is  no 
evidence  of  it  in  embryos  of  the  age  represented  in  Figs. 
4-6.  Cell  division  apparently  takes  place  uniformly  through- 
out the  embryo,  the  result  being  the  building  up  of  a  spherical 
body  (Figs.  4-7).  The  embryo  retains  its  spherical  shape  until 
composed  of  several  hundred  cells.  At  about  this  time  the 
nucellar  tissue  in  the  micropylar  end  of  the  ovule  has  entirely 
broken  down,  so  that  the  embryo  now  lies  in  a  cavity  which  is 
bounded  by  the  inner  integument  of  the  ovule.  Simultaneously 
with  the  division  of  the  embryonal  cell  the  endosperm  nucleus 
divides  and  the  young  embryo  is  soon  surrounded  by  endosperm 


Lyon :  OBSERVATIONS  ON  EMBRYOGENY  OF  NELUMBO.        647 

nuclei  between  which  thin  cell  walls  are  very  early  apparent 
(Figs,  i,  4-7). 

Stage  B.  Monocotyledonous  stage. — The  spherical  embryo 
begins  to  evidence  a  maximum  growth  in  the  horizontal  direc- 
tion, its  greater  dimension  being  diagonal  or  nearly  at  right 
angles  to  the  longer  axis  of  the  ovule.  At  this  time  it  can  per- 
haps best  be  described  as  a  flattened  mass  or  button  of  tissue 
lying  in  the  upper  end  of  the  ovular  cavity.  The  surface  in 
contact  with  the  ovular  wall  conforms  to  the  shape  of  the  latter. 
The  free  surface  is  more  or  less  flattened  and  slightly  inclined 
to  the  plane  of  the  horizontal.  The  plumule  («,  Fig.  8)  now 
arises  as  a  small  protuberance  on  the  inclined  free  surface  in  a 
median  line  near  its  lower  side.  The  axis  of  the  plumule  is 
from  the  first  about  parallel  with  the  axis  of  the  ovule,  and  as 
it  grows  straight  down  into  the  ovular  cavity  it  causes  this  side 
of  the  embryo,  which  we  will  term  the  front  side,  to  become 
flattened.  The  future  cotyledon  is  now  evident  as  a  crescent- 
shaped  mound  of  tissue  (bb,  Fig.  8)  around  the  rear  of  the  em- 
bryo, its  wings  extending  forward  even  with  the  plumule.  This 
stage  culminates  in  an  embryo  as  represented  in  Fig.  8.  The 
endosperm,  during  the  monocotyledonary  stage,  forms  a  co- 
lumnar mass  of  tissue  which  stands  centrally  in  the  cavity,  ex- 
tending from  the  embryo  to  the  persistent  nucellar  tissue  in  the 
lower  portion  of  the  ovule. 

Stage  C.  "Dicotyledonous"  stage. — The  cotyledon  be- 
comes bilobed  through  the  localization  of  growth  at  the  foci,  b 
and  3,  Fig.  8.  From  each  of  these  points  a  cotyledonary  lobe 
grows  rapidly  downward  outside  the  endosperm,  the  tissue  of 
the  nucellus  disorganizing  before  it  (Fig.  10).  In  cross  section 
these  lobes  are  crescent-shaped  (Fig.  16)  and  simultaneously 
with  their  elongation  growth  takes  place  in  both  radial  and  tan- 
gential directions,  each  lobe  at  its  base  growing  forward  around 
the  plumule.  An  idea  of  the  relative  positions  occupied  by  the 
different  structures  may  be  derived  from  Figs.  13-16;  cross 
sections  of  embryos  which  were,  however,  considerably  older 
than  the  one  represented  in  Fig.  9. 

The  growth  of  the  plumule  is  slow  during  this  stage,  it  being 
a  simple  dome-shaped  mound  of  tissue  (Figs.  9,  10  and  18) 
which  comes  to  occupy  a  central  position  through  the  growing 
forward  of  the  cotyledonary  lobes. 

At  about  this  time  the  plerome  first  becomes  apparent  as  a 


648  MINNESOTA    BOTANICAL    STUDIES. 

strand  of  somewhat  smaller  cells  running  from  the  plumule  di- 
rectly through  to  the  base  of  the  embryo  (Figs.  12-18).  From 
this  main  axis,  at  a  point  just  above  its  middle  (Fig.  18),  lateral 
strands  radiate  outward  and  traverse  the  lobes  of  the  cotyledon. 
This  stage  culminates  in  an  embryo  as  represented  in  Fig.  9. 

Stage  D.  Mature  embryo. — This  stage  which  is  reached  with 
the  completion  of  intraseminal  growth  may  be  discussed  under 
the  following  captions  : 

1.  Maturation  of  the  Plumule. — The  first  foliage  leaf  arises 
on  the  axis  of  the  plumule  morphologically  opposite  the  cotyle- 
don (3,  Fig.  14)  (/.  £.,  on  the  front  side  of  the  plumule   axis). 
The   second  foliage  leaf  arises  opposite  the  first  (c,  Fig.  19). 
The   stipules  of    the  first   and  second  leaves   grow  over   and 
around  the  apex  of  the  stem  and  enclose  the  next  two  leaves, 
which  are  also  preformed  in  the  seed  (Fig.  23).     The  lysige- 
nous  cavities  of  the  stem   and  leaf  petioles  very  early  appear, 
as  seen  in  Figs.  19  and  22.     The  structure  of  the  mature  plu- 
mule has  been  many  times  described  and  needs  no  further  treat- 
ment here.      Its   development,  however,  may  be  well  under- 
stood by  a  comparison  of  Figs.  10,  14,  17,  19,  22  and  23. 

2.  Origin    and   Maturation   of  the  Radicle. — The    radicle 
(r,  Fig.    22)  originates   opposite  the  insertion  of  the   plumule 
and  is  but  a  vestigial  structure  not  developing  into  a  functional 
primary  root  upon  the  germination  of  the  seed.     It  only  be- 
comes apparent  at  a  late  stage  in  the  development  of  the  em- 
bryo and  is  usually  completely  enclosed  by  an  outgrowth  of  the 
cotyledonary  tissue.     Fig.  24  shows  a  longitudinal  section  of 
the  hypocotyl  of  a  seedling  which  had  already  developed  sec- 
ondary roots  from  the  epicotyl. 

3.  Maturation  of  the  Cotyledon. — The  edges  of  the  cotyledon- 
ary lobes  soon  meet  through  tangential  growth,  thus  forming  a 
tube  in  which  the  plumule  stands  surrounded  by  the  endosperm. 
The  cells  directly  beneath  the  epidermis   near  the  apex  of  the 
lobes  remain  densely  protoplasmic  and  differentiate  into  a  dis- 
tinct palisade  tissue  (Figs.  20,  21).     This  undoubtedly  forms  a 
nursing  area,  being  in  contact  with  the   lower  portion  of  the 
nucellus  which  persists  for  some  time  and  is  still  evident  in  the 
seed  as  a  thicker,  more  distinct  portion  of  the  testa  (a,  Fig.  n). 
The    lobes    through    more  rapid    elongation    soon    reach  their 
maximum  length  and  then  by  somewhat  slower  tangential  and 
radial  growth  the  embryo  acquires  its  ultimate  spherical  form. 


L\'OH:    OBSERVATIONS  ON  EMBRYOGENY  OF  NELUMBO.  649 

The  integuments  of  the  ovule  keep  pace  with  the  growth  of  the 
embryo  and  form  the  thin  testa  of  the  seed  which  lines  the  thick 
pericarp. 

4.  Fate  of  the  Endosperm. — The  endosperm  reaches  its  max- 
imum growth  early  in   Stage  C  when  it  forms  a  considerable 
mass  of  tissue  lying  between  the  lobes  of  the  cotyledon  (Fig. 
16).     The  cell  walls  are   never  firm  and  offer  little  resistance 
to  the  growing  plumule  which  forces  its  way  into  the  center  of 
the  mass.     The  nuclei  soon  after  begin  to  disappear  and  the 
cells    to   lose  their    definite    outline.     The  resulting    debris    is 
crowded  around  the  plumule  by  the  growth  of  the  cotyledonary 
lobes  and  is  apparent  in  the  seed   as  a  colorless  structureless 
sheath  nearly  or  quite  surrounding  the  plumule. 

5.  The  Embryo  of  the  Seed. — A  earful  study  of  the  embryo 
of  the  seed  without  reference  to  its  development  brings  to  light 
many  conditions  not  to  be  found  in  a  dicotyledonous  seed.     The 
lobes  of  the  cotyledon  are  not  separate  structures,  but  have  a 
common  tissue  at  the  base  of  the  embryo  upon  which  the  plu- 
mule stands  (Figs,  n,  22).     The  sinuses  between  the  lobes  are 
not  of  equal  depth,  the  front  sinus  being  deeper  than  the  rear, 
so  that  the  common   tissue  of  the  lobes   (3,   Fig.    n)   extends 
higher  up  in  the  rear  of  the  plumule  than  it  does  in  front  of  it. 
This  peculiarity   is   even   more   noticeable   in  the  seed  of   N. 
nelumbo  than  in  that  of  N.  lutea  illustrated  in  the  figure,  and  it 
is  so  distinct  that  it  is  remarkable  that  it  has  not  been  described 
before.     The  radicle  is  to  a  greater  or  less  extent  imbedded  in 
tissue  of  cotyledonary  origin,  in  this  respect  conforming  to  well- 
known  monocotyledonous  types. 

THE  EMBRYOGENY  OF  NELUMBO  COMPARED  WITH  THAT 
OF  OTHER  MONOCOTYLEDONS. 

In  its  early  development  the  embryo  of  Nelumbo  is  strikingly 
similar  to  that  of  Pistia  described  by  Hegelmaier  ('74,  68 1- 
686.  pi.  ii .Jig.  4.5-52}.  The  oosperm  of  Pistia,  Hegelmaier 
finds,  does  not  cut  off  a  suspensor  cell  but  by  uniform  divisions 
builds  up  a  spherical  embryo  as  in  Nelumbo.  The  bifurcation 
of  the  cotyledon  is  not  so  inexplicable  a  deviation  from  the  or- 
dinary course  of  development  as  it  may  at  first  appear.  It  is 
rather  to  be  considered  as  a  modification  brought  about  through 
the  adaptation  of  the  embryo  to  the  available  space  within  its  in- 
vestments. 


650  MINNESOTA    BOTANICAL    STUDIES. 

As  the  embryo  departs  from  its  primitive  spherical  form  it  is 
evident  that  its  axis  lies  more  or  less  directly  across  the  ovular 
cavity.  As  the  plumule  develops  superficially  near  one  side  of 
this  flat  expanse  of  tissue  the  remaining  larger  surface  is  to  be 
considered  as  that  of  the  young  cotyledon.  The  cotyledon 
soon  meets  obstruction  to  its  further  growth  in  the  horizontal 
plane  and  hence  turns  down  into  the  ovular  cavity.  Its  later 
bifurcation  is  undoubtedly  the  result  of  its  having  to  adjust  it- 
self to  the  cylindrical  cavity  in  which  it  grows.  That  the 
shape  of  the  cavity  does  to  some  extent  affect  the  shape  of  the 
cotyledon  seems  to  be  evidenced  by  the  fact  that  the  cotyledon- 
ary  lobes  are  at  all  times  closely  appressed  to  their  enclosing  in- 
vestments. The  bifurcation  extends  along  aline  of  mechanical 
stress. 

The  mature  embryo  of  Nelumbo  can  perhaps  best  be  com- 
pared with  those  of  the  grasses,  especially  one  having  but  a 
small  amount  of  cotyledonary  tissue  below  the  junction  of  the 
plumule  and  cotyledon.  The  embryo  of  the  wild  rice,  Ztzania 
aquattca,  as  figured  by  Kennedy  ('99,  pi.  J.  Jig.  22—24)  af- 
fords such  an  example.  It  has  a  long  epicotyl  and  a  very 
short  hypocotyl  which  is  imbedded  in  a  small  amount  of  coty- 
ledonary tissue.  The  cotyledon  grows  around  and  nearly  en- 
closes the  plumule  with  a  uniform  thickness  of  tissue.  In 
Nelumbo  the  conditions  are  almost  exactly  similar,  except  that 
the  cotyledon  in  addition  to  growing  around  the  plumule  has 
become  divided  lengthwise  nearly  to  the  base  into  two  equal 
parts. 

THE  SYSTEMATIC  POSITION  OF  THE  NYMPH^EACE^E. 

As  indicated  above  the  one  character  which  has  led  to  the 
placing  of  the  Nymphagaceae  among  the  Dictotyledons  is  the 
structure  of  the  embryo  in  the  seed.  In  their  other  characters 
they  conform  more  nearly  to  the  Monocotyledons.  The  embryo 
of  Nelumbo  has  been  shown  to  be  monocotyledonous  in  its  de- 
velopment. Although  in  the  other  genera  there  is  present  in 
the  seed  a  functional  endosperm  and  perisperm,  a  careful  ex- 
amination of  the  mature  embryos  shows  them  to  be  in  all  essen- 
tial respects  quite  similar  to  that  of  Nelumbo.  Those  common 
characters,  then,  which  have  previously  united  these  plants  into 
a  family  justify  the  conclusion  that  they  agree  in  embryogeny. 
In  order  to  remove  all  possible  doubt  the  embryogeny  of  a 


Lyon:  OBSERVATIONS  ON  EMBRYOGENY  OF  XELUMBO.         651 

number  of  Nymphaeacece  will  be  studied  during  the  coming 
summer.  The  Nymphceaceae  should  be  classified,  in  the  natural 
system,  in  a  subseries  Nymphaeineas  coordinate  with  the 
Potamogetonineae,  Alismineae  and  Butomineas  in  the  series 
Helobise. 

RECAPITULATION  AND  SUMMARY. 

1.  Jfclumbo  both  in  its   anatomy  and  embryogeny  conforms 
to  the  type  of  the  Monocotyledons. 

2.  The  two  fleshy  bodies  of  the  embryo  arise  through  the  bi- 
furcation of  the  originally  single  cotyledon. 

3.  The  membrane  surrounding  the  plumule  is,  as  conjectured 
by  Wigand,  a  true  endosperm  arising  within  the  embryo-sac. 

4.  The    family  of   the    Nymphasaceae    should    be    classified 
among  Monocotyledons  in  the  series  Helobiae. 

The  writer  wishes  to  express  his  obligations  to  Professor 
Conway  MacMillan  who  has  carefully  followed  the  present  in- 
vestigation, offering  many  suggestions  which  have  been  incor- 
porated in  the  general  discussion. 

LITERATURE. 

The  following  bibliography  includes  the  more  important  works 
dealing  with  the  anatomy,  morphology  or  affinities  of  Nelumbo,  which 
have  been  found  of  value  in  the  present  research. 

1.  Baillon,  H.  (V1)-     Monographic  des  Nymphaeace'es.     Histoire  des 
Plantes,  3:  77-104.      1871. 

2.  Barthelemy,  A.  ('74).     De  la  respiration  et  de  la  circulation  des 
gaz  dans  les  vegetaux.     Ann.  Sci.  Nat.  Bot.  V.  19  :  131-175.     1874. 

3.  Barthelemy,  A.  ('76).     Du  developpement  de  1'embryon  dans  le 
Nelumbium  speciosum  et  de  sa  germination.     Rev.   Sci.  Nat.  5  : 
40-44.      1876. 

4.  Brongniart,  A.  ('27) .     Memoire  stir  la  G£ne>ation  et  le  Developpe- 
ment de  1'Embryon  dans  les  v^ge'taux  phan^rogames.     Chap.   iv. 
Ann.  Sci.  Nat.  12:  225-296.      1827. 

5.  Caspary,  R.  ('78).     Nymphaeaceae  in  v.  Martius.     Fl.  brasiliensis. 
Fasc.  77.  130.      1878. 

6.  Caspary,  R.  ('91).     Nymphaeaceas  in    Die   Natiirl.   Pflanzenfam . 
III.  Teil.  2  Abt.  i-io.     1891. 

7.  Caspary,  R.  ('57).     Flora  717-718,  also  Bot.  Zeit.  791.     1857. 

8.  Clos,  M.  D.   ('91).     Interpretation   des  Parties  Germinatives   du 
Trapa  natans,  de  quelques  Guttiferes  et  des  Nelumbium.     B.  S.  B. 
de  France,  38:  271-276.      1891. 


652  MINNESOTA    BOTANICAL    STUDIES. 

9.  De  Candolle,   A.    ('05).     Sur   la    graine    cles   Nymphcea.     Paris. 
Soc.  Philom.  Bull.  III.  68.     1805. 

10.  De  Candolle,  A.  ('21).     Sur  les  affinites  naturells  de  la  famille 
des  Nympheacees.     Geneve.     Mem.  Soc.  Phys.  I.  208-244.     1821. 

n.  Engelmann,  G.  ('60).     Remarks  on  Ne/um&zum  tuteum.     Trans. 
St.  Louis  Acad.  Sci.  (Proc.)  2.      1860. 

12.  Gaertner,  J.  (1788).     De  Fructibus  et  Seminibus  Plantarum,  I: 

73-74-     i788- 

13.  Gwynne-Vaughan,  D.  T.   ('96,  i).     A  new  case  of  polystely  in 
Dicotyledons.     Ann.  of  Bot.  10  :  289-291.      1896. 

14.  Gwynne-Vaughan,  D.  T.  ('96,  2).     The  arrangement  of  vascular 
bundles  in  certain  Nymphasaceae.     Ann.  of  Bot.  10  :  624-625.    1896. 

15.  Gwynne-Vaughan,  D.  T.   ('97).     On  some  points  in  the  mor- 
phology and  anatomy  of  the  Nymphasaceas.     Trans.  Linn.  Soc.  of 
London,  5  :   287-299.      1897. 

1 6.  Heckel,  E.  ('79)«     Des  poils  et  des  glandes  pileuses  dans  quelques 
genres  de  Nympheacees.     Comp.  Rend.  89  :   758-759.      1879. 

17.  Hegelmaier  ('74)-    Zur  Entwicklungsgeschichte  monokotyledoner 

Keime  nebst  Bemerkungen  iiber  die  Bildung  cler  Samendeckel.    Bot. 
Zeit.  631,  648,  657,  673,  689,  705.      1874. 

i S.  Heritage,  B.  ('95).     Preliminary  notes  on  Nclumbo  lutea.     Bull. 
Torr.  Bot.  Club,  22:    266-271.      1895. 

19.  Jeffrey,  E.  C.     The  morphology  of  the  central   cylinder  in  the 
Angiosperms.     Nymphaeaceae.     24-27. 

20.  Klebs,  G.    ('83).     Beitrage  zur  Morphologic   und  Biologic   der 
Keimung.     Untersuch.  aus  dem  Bot.  Inst.  zu  Tubingen,  I  :    536— 
614.      Nymphaeaceae,  555-556.      1883. 

21.  Kennedy,  P.  B.  ('99).     The  structure  of  the  caryopsis  of  Grasses 
with  reference  to  their  morphology  and  classification.     Bull.  No. 
19,  U.  S.  Dept.  Agric.  Div.  of  Agrostology.      1899. 

22.  Lindley,  J.  ('36).     A  natural  system  of  botany.     Nymphaeaceae, 
10-14.      1836. 

23.  Lyon,  H.  L.   (1901).     Preliminary  note  on  the   embryogeny  of 
Nelumbo.      Science,  13:   470.      1901. 

24.  Mirbel,  C.  F.  B.  ('09).     Observations  anatomiques  ou  physiolo- 
giques  sur  le  Nelumbo  nucifera.     Paris,  Mus.  Nat.  Hist.  Annal.  13  : 
465-481.      1809. 

25.  Mirbel,  C.  F.  B.  ('15).     Siemens  de  Physiologic  Vegetale  et  de 
Botanique,  I  :   59—60. 

26.  Planchon,   J.   E.    ('53).     Etudes   sur   les   Nympheacees.     Ann. 
Sci.  Nat.  Bot.  III.  19:   17-63.      1853. 


Lyon:  OBSERVATIONS  ON  EMBRYOGENY  OF  NELUMBO.         653 

27.  Poiteau,  A.  ('09).     Sur  1'embryon  des  Graminees  des  Cype'race'es, 
et  du  Nelumbo.     Paris,  Mus.  Hist.  Nat.  Annal.  13  :  381-400.     1809. 

28.  Raciborski,  M.   ('94).     Beitriige  zur  Kenntniss  der  Cabombeen 
und  Nymphaeaceen.     Flora,  92-108.      1894. 

29.  Richard,  L.  C.   ('n).     Analyze  botanique  des  embryons   endo- 
rhizes  ou  monocotyledones,  et  particulierement  de  celui  des  Grami- 
n£es.     Paris  Mus.  Hist.  Nat.  Annal.  17  :  223-251,  442-487.     1811. 

30.  Solereder,  H.  ('99).     Systematische  Anatomic  der  Dicotyledonen. 
Nymphaaaceaa.     54-59-      1899. 

31.  Trecul,  A.  ('45).     Recherches  sur  la  structure  des  Nymphaeace'es. 
Tourn.  de  Pharm.  7:   465-467.      1845. 

32.  Trecul,  A.  ('53).     Memoir  sur  la  Formation  des  Feuilles.     Ann. 
Sci.  Nat.  Bot.  III.  20  :   235-314.     1853. 

33.  Trecul,  A.  ('54,  I ).     Etudes anatomiques  et  organageniques  sur  la 
Victoria  regia,  et  anatomic  compared  du  Nelumbium,  du  Nuphar  et 
de  la  Victoria.     Ann.  Sci.  Nat.  Bot.  IV.  I  :   145-172.      1854. 

34.  Trecul,  A.  ('54,  2).     Ve"ge"tation  du  Nelumbium  codophyllum  et 
disposition  anomale  de  ses  Feuilles  et  de  ses  Stipules.     Ann.  Sci. 
Nat.  Bot.  IV.  I  :   291-298.      1854. 

35.  Trecul,  A.  ('54,  3).     Disposition  des  Stipules  et  des  Feuilles  du 
Nelumbium  codophyllum  et  Vegetation  singuli^re  de  cette  Plante. 
B.  S.  B.  de  France,  I  :    18-22,  60-63.      x^54- 

36.  VanTieghem,  P.  ('86,  i).    Surl'apareil  secre"teur  et  les affinite' s  de 
structure  des  Nympheacees.     B.  S.  B.  de  France,  33  :  72—76.     1886. 

37.  Van  Tieghem,  P.    ('86,  2).     Sur  la  croissance  terminate  de   la 
Racine  dans  les  Nymphe'acees.     B.  S.  B.  de  France,  33:   264-265. 
1886. 

38.  Van  Tieghem,  P.,  and  Douliot,  H.  ('88).     Recherches  compara- 
tives sur  1'origine  des  membres  endogenes  dans  les  plantes  vasculaires. 
Ann.   Sci.  Nat.   Bot.  VII.  8:    (Nymphaeaceas)  127-132,  435-43$. 
iSSS. 

39.  Warming,   E.    ('79-'8o).     Forgreningen   og   Bladstillingen  hos 
Sliigten  Nelumbo.     Vidensk.  Meddelelser  fra  den   Naturhistoriske 
Forening  i  Kjobenhavn.  443—455.      1879—80. 

40.  Weberbauer,  A.  ('94).     Beitrage  zur  Samenanatomie  der  Nym- 
plKeaceen.     Bot.  Jahr.  18 :   213-258.      1894. 

41.  Wettstein,  R.  V.  ('88).     Beobachtungen  u'ber  den  Bau  und  die 
Keimung  des  Samens  von   Nelumbo   nucifera   Giirtn.     Verhandl. 
d.  k.  k.  zool.-bot.  Gesellsch.  in  Wien.  38:  41-47.      1888. 

42.  Wigand,  A.  ('71).     Nelumbium  speciosum  W.     Bot.  Zeit.  812- 
826.     1871. 

43.  Wigand-Dennert  ('88).     Nelumbium  speciosum   W.   Eine   mo- 
nographische  Studie.     Bibl.  Bot.  Heft.  n.      1888. 


654  MINNESOTA    BOTANICAL    STUDIES. 


DESCRIPTION  of  PLATE  XLVIII. 

Figure  i.  Longitudinal  section  of  ovule  containing  young  embryo 
and  endosperm  (X5o).  The  nucellar  tissue  about  the  middle  of  the 
embryo-sac  has  broken  down.  The  antipodal  body  is  seen  at  the 
lower  end  of  the  sac. 

2.  Antipodal  body  (xi6o). 

3.  Eight-celled  embryo  (  x  770).      (Outlined  with  camera  lucida.) 
a  =  disorganizing  synei'gid. 

4.  5,  6.   Longitudinal  sections  of  ovules  containing  spherical  embryos 
and  endosperm  (  x  240)  . 

7.  Cross  section  of  ovule  containing  a  spherical  embryo  closely  sur- 
rounded by  endosperm  (  x  240) . 

8.  Front  view  of  an  embryo  at  end  of   stage  B.     a= plumule,  bb-= 
cotyledon.     This  figure   was   reconstructed  from  careful   micrometer 
measurements  of  serial  cross  sections ;   a  proceeding  found  necessary 
in  order  to  get  the  correct  orientation  of  the  embryo. 

9.  Front  view  of  an  embryo  at  end  of  stage    C.     #  =  plumule.     bb 
=  cotyledonary  lobe.     Figure  obtained  in  same  manner  as  Fig.  8. 

10.  Longitudinal    section   of   an  embryo  in   Stage  C  (X5o).      The 
endosperm   is   shrunken   by  reagents   into   the   center  of    the    ovular 
cavity. 

11.  A  fruit  with  one  side  of  the  pericarp  and  a  cotyledonary  lobe 
cutaway  (X2).     #=thicker   portion  of  the  testa.     3=common  base 
of  the  cotyledonary  lobes. 

PLATE  XLIX. 

12.  Cross  section  of  the  hypocotyl  of  an  embryo  just  completing 
Stage  C  ( X  50) . 

13.  Cross   section   of  the   same   embryo  through  base   of  plumule 

(xso). 

14.  Cross  section  of  the   same  embryo   through   apex  of  plumule 
^450).     a=apex  of  stem.     6=first  leaf.      <?=endosperm. 

15.  Cross  section  of  an  embryo,  cutting  plumule  a  short  distance 
above  its  base  (  x  50).     e—  endosperm. 

1 6.  Cross    section    of    same    embryo    through    cotyledonary    lobes 
(X5o).     e=endosperm. 

17.  Longitudinal  section  of  an   embryo  early  in   stage  D    (x65). 
The  fundament  of  the  first  leaf  is  already  evident,  also  the  invagi- 

nation  at  the  base   of  the  embryo,  which   marks  the  position  of  the 
radicle. 


Lyon :   OBSERVATIONS  ox  EMBRYOGENY  OF  NELUMBO.       655 

1 8.  Shows  a  sectional  view  of  the  base  of  an  embryo  in  stage  C. 
(X  190). 

19.  Shows  the  plumule  of  a  young  embryo  in  stage  D  in  longitudi- 
nal section  (  X  50).     a  =  apex  of  stem.     £=first  leaf.     c=  second  leaf. 
c= endosperm.     sl  =stipule  of  first  leaf.     s2  =  stipule  of  second  leaf. 

20.  Vertical   section   through   apex  of   cotyledonary  lobes  showing 
differentiation  of  a  nursing  area  (  x  50) . 

21.  A  portion  of  the  same  under  a  higher  magnification  showing 
the  densely  protoplasmic  palisade  tissue  of  the  nursing  area. 


PLATE  L. 

22.  Longitudinal  section  of  the  plumule  of  the  seed  (XI5).     The 
first  and   second   foliage    leaves   are  cut  off,  their  basal  portions  only 
showing  in  the  figure.      r=  radicle. 

23.  The  apical  bud  in  the  same  section   (  X  50)   showing  the  third 
and   fourth  foliage   leaves   enclosed  by  the   stipules  of  the   first   and 
second.     #=apex  of   stem.     ^=stipule  of   first   leaf.     ^2=stipule  of 
second  leaf. 

24.  Longitudinal  section  of  the  hypocotyl  of  a  seedling  showing  the 
vestigial  radicle  (X5o). 

25.  Cross  section  through  the  epicotyl  of  a  young  seedling  showing 
the  adventitious  roots,  the  first  to  function,  (  x  50) . 


VOL.  II. 


MINNESOTA  BC 


litick 


;AL  STUDIES. 


PART  V. 


10 


6          9 


MINNESOTA  BO 


S^fc&S^iJf 


16 


ICAL  STUDIES. 


PART  V. 


19 


17 


18 


LIX. 


VOL   II.  MINNESOTA  BOTANICAL  STUDIES.  PART  V. 


PLATE  L. 


HINTING    CO..     BOSTON. 


XXXIX.     CONTRIBUTIONS    TO  A  KNOWLEDGE  OF 
THE  LICHENS*OF  MINNESOTA.— VI. 
LICHENS  OF  NORTHWEST- 
ERN MINNESOTA. 


BRUCE  FINK. 


CONSIDERATIONS  OF  DISTRIBUTION  AND  HABITAT. 

The  collections  upon  which  this  paper  is  based  were  made 
during  the  last  half  of  June  and  all  of  July,  1900.  The  region 
traversed  comprises  parts  of  Ottertail,  Beltrami  and  Red  lake 
counties  in  northwestern  Minnesota.  This  region  was  not  sup- 
posed, previous  to  its  study,  to  be  a  rich  field  for  a  lichenist, 
but  because  of  its  relationship  to  other  portions  of  the  state  as  to 
lichen  flora  *  it  seemed  necessary  that  it  should  be  investigated. 
Consequently  the  collection  of  somewhat  more  than  200  species 
as  a  result  of  the  most  extended  collecting  trip  that  I  have  yet 
made  in  Minnesota  fully  met  my  expectations  as  to  probable 
results. 

The  itinerary  was  so  planned  as  to  include  regions  furnish- 
ing the  greatest  possible  variety  of  substrata,  moisture  and 
other  conditions  calculated  to  cause  variations  as  to  lichen  flora. 
Thus  the  most  southern  and  western  points  reached  were  en- 
tirely devoid  of  conifers  and  possessed  an  arboreal  flora  quite 
similar  to  that  of  the  southern  portion  of  the  state.  These  areas 
too  were  near  the  border  between  the  wooded  region  to  the  east 
and  the  prairie  to  the  west.  The  areas  thus  briefly  character- 
ized are  those  about  Battle  lake  and  Thief  River  Falls.  The 
southern  portion  of  the  territory  traversed,  studied  at  Battle 
lake  and  Henning,  gave  a  good  view  of  the  high  morainic  area 
of  the  state  with  its  numerous  rounded  hills  covered  with  granitic 
bowlders  and  calcareous  soil  and  pebbles  and  possessing  a  scant 
growth  of  trees  here  and  there.  This  southern  portion  was 

*  Fink,  B.  Contributions  to  a  knowledge  of  the  Lichens  of  Minnesota. — IV. 
Lichens  of  the  Lake  Superior  Region.  Minnesota  Bot.  Stud.  2:  234.  29  D. 
1899. 

657 


658  MINNESOTA    BOTANICAL    STUDIES. 

also  selected  to  furnish  a  connecting  link  with  the  lichen  flora 
already  studied  farther  south  in  the  state.  To  ascertain  the  re- 
lationship of  northwestern  with  northeastern  Minnesota,  pre- 
viously studied,  as  to  lichen  flora,  two  regions,  Bemidji  and 
Red  lake,  were  chosen,  having  an  abundance  of  conifers  and 
swamps.  These  two  regions  lie  to  the  north  and  east  of  the 
others  previously  named  and  well  within  the  pineries.  It  is  to 
be  regretted  that  none  of  the  area  studied  possessed  rock  expo- 
sures of  any  sort,  similar  either  to  those  in  southern  or  in  north- 
ern and  northeastern  Minnesota.  These  have  since  been 
reached  in  a  study  of  the  northern  boundary  of  the  state  directly 
north  of  the  area  now  under  consideration. 

The  first  area  studied  was  that  about  Battle  lake,  about  150 
miles  northwest  of  Minneapolis,  on  the  border  between  the 
wooded  and  prairie  regions.  The  lichen  habitats  here  are 
granitic  and  lime  bowlders,  earth  and  trees.  The  absence  of 
rock  exposures  here,  as  well  as  elsewhere  in  the  area  studied 
during  the  summer,  detracted  much  from  the  richness  of  the 
flora.  The  trees  in  the  region  about  Battle  lake  are  much  the 
same  as  those  about  Minneapolis  and  in  the  portions  of  south- 
western Minnesota  studied  in  1899.  This  similarity  as  to  ar- 
boreal flora,  as  well  as  the  presence  of  the  granitic  and  calcare- 
ous bowlders  gave  a  lichen  flora  quite  similar  to  that  of  the 
more  southern  regions  named  above.  A  careful  consideration 
of  these  resemblances,  such  as  was  undertaken  for  two  regions 
in  the  second  paper  of  this  series,*  would  of  itself  lead  to  a 
long  and  laborious  article,  and  must  be  omitted  to  give  space 
for  more  important  ecologic  considerations.  A  noticeable  feat- 
ure of  the  lichen  flora  about  Battle  lake  is  that  nearly  all  the  trees 
growing  lichens  are  common  near  the  lakes  and  in  heavy  woods 
back  from  the  lakes,  but  rare  in  woods  back  from  the  lakes  and 
not  heavy.  All  this  is  much  like  the  conditions  about  Minne- 
apolis, as  discussed  in  the  second  paper  of  this  series. f  How- 
ever, turning  to  the  rocks,  they  were  found  literally  covered  with 
lichens  even  up  to  the  very  hill-tops  in  the  morainic  area  south 
of  the  town.  The  whole  number  of  species  of  lichens  collected 
about  Battle  lake  is  in.  Comparing  this  with  the  numbers 


*  Fink,  B.  Contributions  to  a  Knowledge  of  the  Lichens  of  Minnesota. — II. 
Lichens  of  Minneapolis  and  Vicinity.  Minn.  Bot.  Stud,  i  :  703-716.  31  My. 
1897. 

|  Fink,  B.     1.  c.,  705. 


Fink:   LICHENS  OF  NORTHWESTERN  MINNESOTA.  659 

hitherto  recorded  for  various  well-studied  localities  in  the  state 
in  this  series  of  papers,  we  find  the  region  slightly  poorer  in 
lichens  than  any  other  region  in  Minnesota  having  as  much  va- 
riety as  to  substrata. 

The  next  collecting  station  was  at  Henning,  about  35  miles 
to  the  northeast.  This  area  was  selected  especially  for  the 
study  of  the  flora  of  the  Leaf  hills  to  the  south  of  the  town  and 
for  that  of  the  tamarack  and  spruce  swamps,  which  were  not 
found  farther  southwest.  Prominent  among  the  floral  elements 
of  the  swamps  are  species  of  Usnea,  Parmelia  caperata  (L.) 
Ach.,  Parmelia  saxatilis  (L.)  Fr.,  Alectortajubata(I^.)T\^.cVi. 
and  Cetraria  ciliaris  (Ach.)  Tuck.  Other  species  will  be  listed 
toward  the  close  of  these  considerations  in  the  discussion  of 
formations.  Passing  from  the  swamps  to  the  Leaf  hills  three 
miles  away,  all  of  the  above  species  become  rare  or  entirely 
wanting,  though  the  two  Parmelias  are  more  frequent  where 
large  trees  extend,  in  a  few  places,  to  the  tops  of  the  morainic 
hills.  Rinodina  sophodes  (Ach.)  Nyl.  and  Arthonia  lecideella 
Nyl.,  become  abundant  on  the  shrubs  of  oak,  poplar  and  birch 
in  the  hills.  These  hills  probably  reach  an  elevation  of  1,800 
feet  south  of  Vining  and  form  the  hightest  morainic  area  in  the 
state.  The  drift  bowlders  and  pebbles  of  granite  and  limestone 
extend  to  the  very  summits  of  the  hills  and  furnish  an  excellent 
field  for  the  study  of  lichen  formations,  especially  of  granitic 
bowlders.  The  calcareous  matter  of  the  drift  has  been  ground 
fine  as  powder  or  left  as  small  pebbles.  The  soil  contains 
enough  lime  so  that  the  lichen  formation  characteristic  of  cal- 
careous earth  is  well  developed,  and  the  limy  pebbles  also 
support  a  well  developed  calcareous  rock-lichen  formation. 
Black  peak  about  four  miles  south  of  Vining  is  an  especially 
good  place  for  the  study  of  the  last  two  formations,  which  will 
receive  careful  consideration  later  in  this  paper.  It  may  be 
noted  in  passing  that  the  lichen  flora  of  this  second  region  is  a 
much  richer  one  than  that  first  studied  near  the  border  of  the 
wooded  region,  where  the  morainic  hills  are  less  developed, 
where  trees  are  less  numerous  and  of  fewer  species  and  where 
the  tamarack  swamps  are  wanting.  In  the  region  about  Hen- 
ning 140  species  and  varieties  were  collected  in  about  the  same 
time  as  was  required  to  find  the  in  at  Battle  lake. 

Passing  on  to  the  next  area  studied  at  Bemidji  75  miles 
north  of  Henning  and  well  within  the  pineries  where  there  is 


660  MINNESOTA    BOTANICAL    STUDIES. 

more  variety  as  to  trees,  154  species  were  collected,  though  the 
calcareous  formations  were  entirely  absent  and  the  granitic  only 
poorly  developed.  The  disadvantage  due  to  absence  of  granitic 
and  calcareous  lichen  formations  seemed  to  be  more  than  offset 
by  the  unusual  richness  of  the  formations  of  the  tamarack  and 
the  cedar  swamps  and  the  earth  under  the  pines.  As  this  \vas 
my  first  opportunity  for  a  careful  study  of  these  three  forma- 
tions, and  the  tree  formations  were  also  especially  rich  as  well 
there,  somewhat  more  time  was  taken  for  the  collecting  at  Be- 
midji  than  elsewhere.  However,  notwithstanding  the  scarcity 
of  any  kind  of  bowlders  and  the  absence  of  rock  exposures, 
the  Bemidji  region  may  be  regarded  as  one  of  the  richest  lichen 
floral  areas  in  Minnesota,  while  Henning  with  its  greater  variety 
of  substrata  and  favorable  conditions  is  scarcely  inferior. 

From  Bemidji  I  passed  to  Thief  River  Falls,  about  80  miles 
to  the  northwest.  Here  I  found  a  territory  composed  for  the  most 
part  of  low  flat  prairie,  but  with  good  woods  along  the  banks 
of  Red  lake  and  Thief  rivers.  The  trees  are  mostly  birches, 
poplars,  oaks  and  elms,  forming  a  monotonous  arboreal  flora 
by  no  means  favorable  to  the  production  of  a  large  number  of 
lichens.  Inspection  of  the  list  of  species  and  varieties  recorded 
for  this  region  will  show  that  only  43  lichens  were  found  on 
these  trees  to  compare  with  more  than  twice  as  many  on  trees 
at  Bemidji,  and  that  these  43  are  in  general  the  most  common 
of  Minnesota  lichens  growing  on  trees.  Marshes  are  frequent, 
and  devoid  of  trees ;  or  having  the  same  species  as  grow  on  the 
higher  ground,  sustain  no  peculiar  lichen  species.  Both  granitic 
and  lime  bowlders  are  frequently  seen  in  the  region,  but  they 
are  almost  totally  bare  of  any  sort  of  plant  life.  Doubtless  this 
is  due  partly  to  fires  which  frequently  run  over  the  prairies.  In 
places  pastured  for  several  years  so  that  fires  have  not  occurred, 
lichens  are  beginning  to  take  possession  of  the  rocks.  Yet  it  is 
difficult  to  explain  the  absence  of  lichens  along  high  bowlder, 
strewn  river  banks,  as  in  certain  Localities  toward  St.  Hilaire, 
on  any  supposition.  It  will  be  very  interesting  to  note  the  in- 
crease of  lichen  flora  on  the  rocks  of  this  region  as  the  country 
becomes  settled  more  densely  and  fires  are  kept  out.  In  order 
that  this  may  be  done,  I  record  the  few  lichens  now  occurring 
rarely  on  these  rocks. 

Rinodina  oreina  (VILL.)  MASS. 

Lecanora  varia  (EHRH.)  NYL. 


Fink:    LICHENS  OF  NORTHWESTERN  MINNESOTA.  661 

Lecanora  varia  (EHRH.)  NYL.  var.  polytropa  NYL. 

Placodium  murorum  (HOFFM.)  DC. 

Placodium  elegans  (LINK)  DC. 

Physcia  caesia  (HOFFM.)  NYL. 

Parmelia  conspersa  (EHRH.)  ACH. 

The  earth  lichen  flora  is  as  poorly  developed  as  that  of  the 
trees  and  rocks,  and  the  whole  known  lichen  population  of  the 
area  comprises  only  78  species  and  varieties  and  is  the  most 
scanty  yet  studied  in  the  state,  except  that  at  Pipestone,  where 
trees  are  almost  wholly  absent. 

The  last  collecting  ground  was  at  Red  lake,  some  65  miles 
east  and  somewhat  south  of  Thief  River  Falls.  This  area  is 
about  36  miles  north  of  the  one  previously  studied  at  Bemidji 
and  has  a  lichen  community  very  similar.  Here  the  only 
bowlders  that  gave  any  noteworthy  results  were  those  along  the 
lake  shore,  and  the  lichens  on  them  were,  all  but  three  or  four, 
of  the  same  species  as  those  growing  upon  the  adjacent  trees. 
With  this  dearth  of  rock  lichens  the  territory,  probably  not 
quite  so  thoroughly  studied  as  the  one  to  the  south  about 
Bemidji,  gave  only  120  lichen  species  and  varieties. 

Compared  with  other  portions  of  the  state  of  equal  size, 
whose  lichen  floras  have  been  investigated,  this  one  is  some- 
what the  poorest  in  lichens.  The  number  collected  is  little 
larger  than  that  found  in  southwestern  Minnesota,  but  should 
be  considerably  larger,  as  fully  one-third  more  time  was  taken 
for  the  collecting.  The  Lake  Superior  region  gave  258  lichen 
forms  in  about  the  same  time  as  was  spent  in  making  the  col- 
lections in  northwestern  Minnesota.  However,  this  is  what 
would  be  expected  since  the  former  area  is  more  diversified  as 
to  climate,  the  portion  near  the  lake  having  many  arctic  and 
subarctic  species,  while  the  northern  and  western  portions 
yielded  essentially  the  same  species  as  the  region  now  under 
consideration.  Then  too  the  absence  of  the  great  exposures  of 
igneous  rocks  of  the  Superior  region  has  already  been  noted 
for  the  present  one,  in  which  only  58  species  and  varieties  of 
lichens,  or  about  28  per  cent,  of  the  whole  lichen  flora,  were 
found  on  rocks,  whereas  nearly  50  per  cent,  of  these  plants  in 
the  former  region  were  collected  on  the  rocks.  The  occurrence 
of  about  three-fourths  of  the  entire  number  of  lichens  of  the 
whole  area  under  consideration  in  one  small  area  about  Bemidji 
demonstrates  that  little  of  the  difference  in  the  composition  of 


662  MINNESOTA    BOTANICAL    STUDIES. 

the  flora  of  various  portions  of  the  territory  is  due  to  edaphic 
causes.  What  variety  exists  is  due  for  the  most  part  to  the 
partial  extension  of  the  conifers,  tamarack  swamps  and  calcar- 
eous soil,  bowlders  and  pebbles  over  the  region. 

The  comparative  richness  of  different  collecting  grounds  as 
to  lichen  species  has  been  noted  in  passing.  It  may  be  added 
that  the  number  of  species  occurring  in  each  place  and  not  else- 
where found  bears  some  relation  to  the  whole  number  of  species 
found  in  each  area  studied.  Thus  Battle  lake  with  a  total  of 
in  collected  has  7  .not  collected  elsewhere.  The  numbers  for 
other  collecting  grounds  are :  Henning  140  and  18,  Bemidji 
154  and  31,  Thief  River  Falls  78  and  2  and  Red  lake  120  and 
14.  Comparison  of  the  numbers  shows,  as  would  be  expected, 
that  the  number  of  rare  lichens  collected  in  the  best  collecting 
grounds  is  in  much  larger  proportion  to  the  total  number  col- 
lected than  in  the  areas  less  favored  as  to  lichen  flora,  where 
one  finds  only  the  commonest  species. 

Of  the  41  lichen  species  recorded  in  the  fourth  paper  of  this 
series  as  arctic  or  subarctic,*  the  following  seven  occur  in  the 
region  considered  in  this  paper,  while  no  new  northern  forms 
were  discovered. 

Ramalina  pusilla  (PREV.)  TUCK. 
Usnea  cavernosa  TUCK. 
Stereocaulon  paschale  (L.)  FR. 
Cladonia  deformis  (L.)  HOFFM. 
Cladonia  digitata  (L.)  HOFFM. 
Biatora  leucophaea  (FLK.)  TUCK. 
Buellia  petraea  (FLOT.,  KBR.)  TUCK. 

The  other  species  not  new  to  Minnesota  are  in  general  those 
found  farther  south  in  the  state.  Thus  the  prediction,  as  to  re- 
semblance of  the  present  flora  to  that  farther  south  in  Minne- 
sota, made  in  the  fourth  paper  of  this  series, f  seems  to  be  fairly 
well  established,  though  the  extreme  northern  boundary  of  the 
state  west  of  the  Snowbank  lake  area  remains  to  be  considered 
in  the  next  paper.  Of  the  genera  having  northern  species  in 
northeastern  Minnesota,  Solorina^Heterothecium,  Bceomyces^nd 
Umbilicaria  were  not  found  in  the  area  now  under  consideration, 
nor  was  there  found  more  than  a  single  genus,  Melaspilea,  new 
to  the  state.  Moreover,  the  paucity  of  lichens  in  northwestern 

*  Fink,  B.     1.  c.,  227-232. 
fFink,  B.     1.  c.,  233-234. 


Fink:    LICHENS  OF  NORTHWESTERN  MINNESOTA.  663 

as  compared  with  northeastern  Minnesota  extends  to  genera  as 
well  as  to  species,  so  that  36  genera  were  found  in  the  former 
area  and  39  in  the  latter.  This  would  be  expected  when  we 
notice  that  all  the  seven  northern  species  recorded  above  belong 
to  genera  occurring  in  the  southern  half  of  Minnesota,  while 
the  four  genera  named  above  as  not  occurring  in  northwestern 
Minnesota  also  have  no  representatives  in  the  southern  half  of 
the  state,  being  for  the  most  part  the  most  strictly  northern  gen- 
era thus  far  found  in  Minnesota. 

The  summer's  collecting  brought  to  light  48  species  and  va- 
rieties new  to  the  state,  of  which  9  are  new  to  North  America 
and  3  new,  while  another  3  are  yet  undetermined.  The  dis- 
covery of  so  large  a  proportion  of  new  material  nearly  one- 
fourth  of  all  species  collected,  after  other  parts  of  the  state  had 
been  for  the  most  part,  well  studied,  is  somewhat  of  a  surprise, 
and  seems  to  indicate  that  there  is  yet  a  good  number  of  lichens 
to  be  found  in  Minnesota.  As  to  forms  new  to  North  America, 
this  paper  adds  a  larger  number  than  all  the  previous  lists  pub- 
lished for  the  state. 

The  genus  Calicium  deserves  special  mention  because  of  the 
interesting  facts  disclosed  when  the  material  collected  was  care- 
fully studied.  Previously  nine  species  and  varieties  had  been 
recorded  for  the  state,  and  though  the  genus  is  not  an  arctic  or 
subarctic  one,  I  had  not  expected  to  find  it  well  represented  in 
territory  otherwise  closely  related  to  the  southern  half  of  Minne- 
sota as  to  lichen  population.  Consequently,  I  was  not  a  little 
surprised  to  find  ten  species  in  my  collection,  one  more  than  had 
hitherto  been  reported  for  the  state,  and  yet  more  when  I  found 
that  six  of  the  species  were  new  to  the  state,  thus  raising  the 
number  of  Caliciums  in  Minnesota  to  15  species  and  varieties. 
We  now  have  found  within  the  state  about  half  of  the  forms  of 
the  genus  recorded  for  North  America,  and  Calicium  can  no 
longer  be  regarded  as  a  genus  peculiar  to  the  Atlantic  region. 
The  cause  of  the  extension  of  members  of  the  genus  into  a  re- 
gion closely  related  with  southern  Minnesota  as  to  lichen  flora 
is  to  be  found  in  the  circumstance  that  the  Caliciums  seem  to 
follow  the  conifers  regardless  of  slight  climatic  changes.  While 
some  other  genera  furnish  each  a  few  species  new  to  the  state, 
or  not  found  further  south  in  Minnesota,  there  is  nothing  espe- 
cially noteworthy  about  the  distribution  of  any  of  them,  as  the 
species  are  in  general  such  as  could  be  expected  to  occur  farther 
south,  and  may  have  been  overlooked. 


664  MINNESOTA    BOTANICAL    STUDIES. 

It  now  remains  to  follow  out  the  ecologic  study  begun  in  the 
fifth  paper  of  this  series,*  pursuing  the  general  plan  of  study 
introduced  there.  First  of  all  I  shall  consider  some  of  the  for- 
mations therein  recorded  and  draw  some  comparisons.  As  in 
the  former  paper  I  shall  consider  especially  the  more  common 
lichens,  which  give  character  to  the  flora  and  lichen  formations 
of  the  state.  Beginning  with  the  lichen  formation  of  the  ex- 
posed granitic  bowlders  about  Battle  Lake,  the  formation  may 
be  designated  as  follows  : 

LECANORA  LICHEN  FORMATION  OF  EXPOSED  GRANITIC 
BOWLDERS  (BATTLE  LAKE). 

Physcia  stellaris  (L.)  TUCK.  var.  apiola  NYL.,  C. 

Physcia  caesia  (HOFFM.)  NYL.,  CX. 

Placodium  elegans  (LINK.)  DC.,  C. 

Placodium  cinnabarrinum  (Acn.)  ANZ.,  C. 

Placodium  aurantiacrm  (LIGHTF.)  NAEG.  and  HEPP.,  B. 

Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP.,  B. 

Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP.  var.  sideritis 
TUCK.,  CX. 

Placodium  vitellinum  (EHRH.)  NAEG.  and  HEPP.,  CX. 

Lecanora  rubina  (VILL.)  ACH.,  CX. 

Lecanora  muralis  (SCHREB.)  ScHAER.var  saxicola  SCHAER.,  C. 

Lecanora  hageni  ACH.,  C. 

Lecanora  varia  (EHRH.)  NYL.,  B. 

Lecanora  varia  (EHRH.)  NYL.  var.  polytropa  NYL.,  B. 

Lecanora  cinerea  (L.)  SOMMERF.,  CX. 

Lecanora  calcarea  (L.)  SOMMERF.  var.  contorta  FR.,  C. 

Lecanora  xanthophana  NYL.,  CX. 

Lecanora  cervina  (PERS.)  NYL.,  C. 

Lecanora  fuscata  (SCHRAD.)  TH.  FR.,  C. 

Rinodina  oreina  (Acn.)  MASS.,  C. 

Rinodina  sophodes  (Acn.)  NYL.,  CX. 

Rinodina  sophodes  (AcH.)  NYL.  var.  exigua  FR.,  B. 

Buellia  pullata  TUCK.,  C. 

Buellia  petraea  (FLOT.,  KBR.)  TUCK.,  CX. 

Buellia  petraea  (FLOT.,  KBR.)  TUCK.  var.  montagnaei 
TUCK.,  C. 

*  Pink,  B.  Contributions  to  a  Knowledge  of  the  Lichens  of  Minnesota. — V. 
Lichens  of  the  Minnesota  Valley  and  Southwestern  Minnesota.  Minn.  Bot. 
Stud.  2  :  283-308.  29  D.  1899. 


Fink :   LICHENS  OF  NORTHWESTERN  MINNESOTA.  665 

This  formation  may  be  compared  est  with  the  si  milar  one 
upon  the  exposed  granite  at  Granite  Falls.*  As  might  be  ex- 
pected the  Battle  lake  formation  confined  to  the  bowlders 
suffers  somewhat  in  comparison  with  that  at  Granite  Falls, 
which  is  developed  upon  extensive  exposures  of  granite.  The 
number  of  lichens  in  the  formation  at  the  former  place  is  24 
and  the  number  at  the  latter  31.  Species  common  to  the  two 
similar  formations  I  have  marked  (C),  and  those  found  at 
Battle  lake  only  are  marked  (B).  Of  the  ten  species  found  at 
Granite  Falls  and  not  at  Battle  lake,  Parmelia  cons-persa 
(Ehrh.)  Ach.  deserves  special  mention  as  being  unaccountably 
rare  in  the  region  covered  by  this  paper.  The  entire  absence 
of  another  of  the  ten,  Biatora  rtifontgraTuck.,  from  the  region 
is  quite  as  remarkable.  The  other  eight  are  lichens  either  not 
widely  distributed  in  the  state,  or  not  common  on  granitic  rocks. 
Eight  of  the  species  of  the  formation  recorded  above  have  oc- 
curred in  all  of  the  four  localities  in  the  state  in  which  similar 
formations  have  been  recorded.  These  may  be  considered  the 
most  constant  elements  of  such  formations  in  Minnesota,  and  I 
have  marked  them  (X).  The  formation  is  also  well  developed 
in  the  Leaf  hills,  but  nothing  would  be  gained  by  recording  it. 

As  to  the  general  character  of  the  lichens  of  the  formation, 
all  but  the  first  three  and  the  first  Lecanora  are  strictly  crusta- 
ceous,  and  this  last  plant,  as  well  as  the  foliaceous  Placodium, 
is  nearly  as  closely  adnate  to  the  substratum  as  the  crustaceous 
forms.  The  less  adnate  foliaceous  species  have  a  well-devel- 
oped cellular  cortex  on  all  sides  for  support  and  for  protection 
against  too  rapid  evaporation  of  moisture  in  their  exposed 
and  dry  habitat.  Of  the  crustaceous  species,  the  Placodiums 
have  good  upper  and  some  of  them  a  lower  cortex,  the  Lecan- 
oras  have  either  an  upper  only  or  none,  while  the  Rinodinas, 
except  possibly  the  first,  have  no  cortical  layers ;  and  the  last 
statement  applies  to  the  Buellias  as  well.  The  forms,  having 
no  cortex  or  a  poorly-developed  one  and  growing  in  such  a  dry, 
exposed  habitat,  have  very  small  thalli.  A  large  foliaceous 
lichen  with  no  cortex,  as  a  Collema  for  instance,  could  hardly 
exist  in  the  present  formation.  Finally  the  lichens  of  the  for- 
mations are  a  few  foliaceous  species  with  well-developed  cortex 
above  and  below,  but  still  quite  closely  adnate,  and  a  much 
larger  number  with  no  cortex  or  an  upper  one  only,  but  having 

*Fink,  B.     1.  c.,  286,  287. 


666  MINNESOTA    BOTANICAL    STUDIES. 

very  small  thalli,  and  though  epilithic  yet  very  closely  adnate, 
so  that  moisture  easily  passes  into  them  from  the  rocky  sub- 
stratum below. 

Next  may   naturally  follow  the  mixed    lichen   formation  of 
shaded  granitic  bowlders. 

MIXED   LICHEN   FORMATION    OF  SHADED   GRANITIC    BOWLDERS 

(RED  LAKE). 

A.  Probably  naturally  belonging  to  the  rocks. 

Biatora  inundata  FR. 
Verrucaria  nigrescens  PERS. 
Verrucaria  viridula  ACH. 
Verrucaria  muralis  ACH. 

B.  Near  trees  and  probably  migrated  from  them. 

Theloschistes  lychneus  (NYL.)  TUCK.,  C. 

Parmelia  borreri  TURN.,  C. 

Parmelia  saxatilis  (L.)  FR.,  C. 

Parmelia  olivacea  (L.)  ACH. 

Parmelia  caperata  (L.)  ACH.,  C. 

Physcia  speciosa  (WULF.,  ACH.)  NYL.,  C. 

Physcia  pulverulenta  (SCHREB.)  NYL.,  C. 

Physcia  stellaris  (L.)  TUCK. 

Physcia  stellaris  (L.)  TUCK.  var.  apiola  NYL. 

Physcia  astroidea  (FR.)  NYL. 

Physcia  hispida  (SCHREB.,  FR.)  TUCK. 

Physcia  obscura  (EHRH.)  NYL.,  C. 

Of  the  12  species  of  the  formation  probably  having  migrated 
from  trees  near  by,  seven  marked  (C)  are  common  to  all  of 
the  six  similar  formations  studied  in  the  state,*  but  none  of  the 
four  elements  naturally  belonging  to  the  rocks  are  common  to 
the  similar  formations. 

Comparing  these  lichens  with  those  of  the  exposed  rock 
formation  above,  we  find  an  entirely  different  type  of  thallus 
to  prevail,  viz.,  the  foliaceous  type  of  the  Parmelias  and 
Physcias  with  thalli  having  well  developed  cortical  layers. 
And  as  would  be  expected  in  an  ombrophytic  lichen  formation 
having  such  thalli,  the  plants  are  not  so  closely  adnate  to  the 
substratum  as  those  of  the  exposed  rock  formation,  but  are 

*Fink,  B.     1.  c.,  290-293. 


Fink:   LICHENS  OF  NORTHWESTERN  MINNESOTA.  667 

more  loosely  attached  by  rhizoids.  However,  those  undoubtedly 
belonging  to  the  rocks  have  poorly  developed  thalli  with  no 
cortex  or  an  upper  pseudo-cortex,  and  Verrucaria  muralis  Ach., 
an  intruder  usually  growing  upon  calcareous  rocks  in  exposed 
places,  is  essentially  hypolithic.  That  the  three  forms  having 
poorly  developed  and  epilithic  thalli  should  seek  ombrophytic 
associations  is,  of  course,  natural  enough. 

The  Cladonia-Pclligera  formation  of  shaded  earth  was  found 
remarkably  well  developed  under  the  pines  at  Bemidji.  The 
table  of  species  below  shows  20  forms.  The  formation  here, 
though  containing  the  same  genera  as  the  similar  one  farther 
south,*  is  twice  as  well  developed,  being  especially  rich  in 
Cladonias,  which  nourish  under  the  pines.  I  have  marked  (C) 
the  five  species  common  to  the  similar  formations  noted  for  the 
state,  as  being  the  most  constant  elements  of  such  formations, 
at  least  in  Minnesota.  I  shall  now  record  the  formation  and 
follow  with  a  discussion  of  structure  of  the  lichens  composing  it. 

CLADONIA-PELTIGERA  LICHEN  FORMATION  OF  SHADED  EARTH 

(BEMIDJI). 

Peltigera  horizontalis  (L.)  HOFFM. 

Peltigera  canina  (L.)  HOFFM.,  C. 

Peltigera  canina  (L.)  HOFFM.  var.  spuria  ACH. 

Peltigera  canina  (L.)  HOFFM.  var.  sorediata  SCHAER.,  C. 

Collema  pulposum  (BERNH.)  NYL.,  C. 

Collema  limosum  ACH. 

Collema  crispum  BORR. 

Cladonia  cariosa  (Acn.)  SPRENG. 

Cladonia  pyxidata  (L.)  FR.,  C. 

Cladonia  degenerans  FLK. 

Cladonia  gracilis  (L.)  NYL.,  C. 

Cladonia  gracilis  (L.)  NYL.  var.  symphycarpia  TUCK. 

Cladonia  gracilis  (L.)  NYL.  var.  verticillata  FR.,  C. 

Cladonia  gracilis  (L.)  NYL.  var.  elongata  FR. 

Cladonia  gracilis  (L.)  NYL.  var.  hybrida  SCHAER. 

Cladonia  cornuta  (L.)  FR. 

Cladonia  furcata  (Huos.)  FR. 

Cladonia  rangiferina  (L.)  HOFFM. 

Cladonia  rangiferina  (L.)  HOFFM.  var.  alpestris  L. 

Cladonia  uncialis  (L.)  FR. 

*Fiuk,  B.     1.  c.,  294-295. 


668  MINNESOTA    BOTANICAL    STUDIES. 

Comparing  the  lichens  of  this  formation  with  those  of  the 
shaded  rock  formation  above,  we  find  that,  as  a  result  of  the 
more  moist  habitat  of  the  forms,  growing  on  earth  in  shade  and 
usually  on  an  abundance  of  decaying  vegetable  remains  which 
hold  moisture,  the  formation  consists  of  plant  individuals  even 
less  closely  attached  to  the  substratum,  except  perhaps  the 
Collemas,  which  have  no  cortex  and  which,  therefore,  even  in 
their  shaded  situation,  remain  close  to  the  substratum  to  absorb 
the  moisture  which  is  rapidly  evaporated  from  their  non-cellular 
surfaces.  The  Peltigeras  with  a  well  developed  upper  cortex 
hold  moisture  better  and  rise  somewhat  higher.  The  podetia  of 
the  Cladonias  have  a  surrounding  pseudo-cortex  of  densely 
interwoven  hyphae,  which  serves  for  protection  ;•  against  too 
rapid  evaporation  of  moisture  as  well  as  for  mechanical  sup- 
port. Hence  the  Cladonias  rise  vertically  and  are  especially 
numerous  as  to  species  and  individuals  in  this  moist  and  shaded 
formation,  as  well  as  unusually  luxuriant.  Thus  it  appears 
that  we  have  in  this  formation,  as  in  those  already  considered 
plants  adapted  in  very  different  ways  to  the  environment. 

The  remarkable  constancy  of  occurrence  of  certain  floral 
elements  in  certain  environments  can  scarcely  be  better  illus- 
trated than  by  comparing  the  calcareous  earth  lichen  formation 
given  below  with  similar  ones  recorded  in  the  fifth  paper  of  this 
series  for  Granite  Falls,  Minnesota,  and  Fayette,  Iowa.* 

BIATORA    DECIPIENS     LICHEN    FORMATION~OF    EX-POSED    CALCA- 
REOUS EARTH  (LEAF  HILLS). 

Heppia  despreauxii  (MONT.)  TUCK.,  C. 

Urceolaria  scruposa  (L.)  NYL.,  C. 

Biatora  decipiens  (EHRH.)  FR.,  C. 

Biatora  decipiens  (EHRH.)  FR.  var.  delabata  AUCT.,  C. 

Biatora  muscorum  (Sw.)  TUCK.,  C. 

Endocarpon  hepaticum  ACH.,  C. 

All  of  the  six  floral  elements  recorded  in  the  present  forma- 
tion and  marked  (C)  are  also  listed  in  the  formation  at  Granite 
Falls  and  Fayette,  and  each  of  these  last  two  contains  a  single 
rare  species  not  discovered  in  the  Leaf  hills  formation.  As  in 
the  localities  previously  studied,  the  formation  in  the  hills  is  best 
developed  on  the  hill-sides  where  the  plants  are  washed  with 
the  lime-impregnated  water  which  flows  down  the  slope  during 

*Fink,  B.     1.  c.,  295-296. 


Fink:    LICHENS  OF  NORTHWESTERN  MINNESOTA.  669 

rains,  and  it  may  be  regarded  as  well  established  that  such  a 
sloping  surface  is  best  adapted  to  the  development  of  the  forma- 
tion. 

As  to  structural  adaptations  to  an  exposed  and  usually  dry 
habitat,  the  lichens  of  the  formation  all  have  small  thalli  and 
are  closely  adnate.  The  rudimentary  thallus  of  Biatora  mus- 
corum  (Sw.)  Tuck,  has  no  cortex,  and  the  somewhat  better  de- 
veloped one  of  the  Urceolaria  has  only  a  poorly  developed 
pseudo-cortex  above.  Of  the  larger  and  better  developed  thalli, 
that  of  Biatora  decipiens  (Ehrh.)  Fr.  has  a  very  heavy  cellular 
cortex  above,  the  Endocarpon  has  a  well-developed  cortex  on 
all  sides  while  the  Heppia  is  cellular  throughout.  In  these 
larger  thalli  growing  in  exposed  dry  places,  the  cellular  areas 
serve  of  course  not  only  for  support,  but  also  for  protection  of 
the  algge  within  and  against  excessive  evaporation ;  nor  must  it 
be  supposed  that  any  of  these  thalli  are  large  for  a  really  large 
thallus  is  seldon  seen  in  such  a  formation,  and  if  present  at  all, 
should  be  considered  an  accident  in  plant  distribution  not  to  be 
recorded.  The  four  better  developed  thalli  are  then  only  large 
in  comparison  with  the  other  two  and  average  about  3  to  6  mm. 
in  diameter.  The  two  rudimentary  thalli  of  the  formation  are 
able  to  persist  because  very  small  and  closely  adnate. 

Comparing  next  the  lichen  formation  of  the  calcareous  peb- 
bles of  the  same  area  with  the  same  two  for  the  Minnesota  and 
Iowa  localities  used  above,*  we  find  that  the  adding  of  another 
Minnesota  locality  still  leaves  the  same  five  species  common  (C) 
to  such  formations  for  the  two  states.  This  third  formation  of 
the  kind  recorded  below,  establishes  a  general  resemblance  of 
such  formations  in  widely  separate  localities. 

LECANORA    CALCAREA    CONTORTA    LICHEN    FORMATION    OF     EX- 
POSED   LIMESTONE    (LEAF    HILLS). 

Placodium  vitellinum(EHRH.)NAEG.  and  HEPP.  var.  aurellum 
ACH.,  C. 

Lecanora  muralis  (SCHREB.)  SCHAER.  var.  versicolor  FR. 

Lecanora  subfusca  (L.)  ACH. 

Lecanora  calcarea  (L.)  SOMMERF.  var.  contorta  FR.,  C. 

Lecanora  privigna  (Acn.)  NYL.,  C. 

Lecanora  privigna  (Acn.)  NYL.  var.  pruinosa  AUCT. 

Rinodina  bischoffii  (HEPP.)  KBR. 

*Fink,  B.     1.  c.,  297 


670  MINNESOTA    BOTANICAL    STUDIES. 

Endocarpon  pusillum  HEDW.,  C. 

Verrucaria  nigrescens  PERS. 

Verrucaria  muralis  ACH.,  C. 

The  lichens  of  the  above  formation  have  small  thalli,  closely 
adnate  or  even  more  or  less  strictly  hypolithic.  All  except  the 
last  Verrucaria,  which  is  hypolithic,  and  Lecanora  -pri-vigna 
(Ach.)  Nyl.,  which  has  a  very  rudimentary  and  evanescent 
thallus  have  more  or  less  of  a  cellular  or  pseudocellular  cortex 
above.  None  except  the  first  Lecanora,  which  has  the  largest 
thallus  in  the  formation,  showed  any  indication  of  such  structure 
below.  The  upper  cortex  gives  these  better  developed  but  still 
small  thalli  sufficient  protection  against  evaporation  and  adapts 
them  to  their  dry  station. 

The  lichen  formation  for  trees  with  rough  and  smooth  bark 
at  Bemidji  are  in  general  quite  like  the  corresponding  ones 
recorded  for  Mankato  and  Granite  Falls,*  but  are  rather  richer 
in  species.  I  record  them  below  as  they  may  be  of  use  in  the 
further  study  of  the  lichens  of  northern  Minnesota. 

PARMELIA  LICHEN  FORMATION  OF    TREES  WITH  ROUGH  BARK 

(BEMIDJI). 

Ramalina  calicaris  (L.)  FR.  var.  fastigiata  FR. 

Ramalina  calicaris  (L.)  FR.  var.  fraxinea  FR. 

Usnea  barbata  (L.)  FR.  var.  florida  FR. 

Theloschistes  chrysopthalmus  (L.)  NORM. 

Theloschistes  polycarpus  (EHRH.)  TUCK. 

Theloschistes  lychneus  (NYL.)  TUCK. 

Theloschistes  concolor  (DICKS.)  TUCK. 

Theloschistes  concolor  (DICKS.)  TUCK.  var.  effusa  TUCK. 

Parmelia  crinita  ACH. 

Parmelia  borreri  TURN. 

Parmelia  tiliacea  (HOFFM.  )  FLK. 

Parmelia  saxatilis  (L.)  FR. 

Parmelia  saxatilis  (L.)  FR.  var.  sulcata  NYL. 

Parmelia  olivacea  (L.)  ACH. 

Parmelia  olivacea  (L.)  ACH.  var.  sorediata  (Acn.)  NYL. 

Parmelia  caperata  (L.)  ACH. 

Physcia  speciosa  (WULF.,  ACH.)  NYL. 

Physcia  hypoleuca  (MUHL.)  TUCK. 

*Fink,  B.     1.  c.,  302-305. 


Pink:    LICHENS    OF    NORTHWESTERN    MINNESOTA.  671 

Physcia  pulverulenta  (SCHREB.)  NYL. 
Physcia  stellaris  (L.)  TUCK. 
Physcia  hispida  (SCHREB.,  FR.)  TUCK. 
Physcia  obscura  (EHRH.)  NYL. 
Collema  pycnocarpum  NYL. 
Collema  flaccidum  ACH. 
Collema  nigrescens  (HUDS.)  ACH. 
Leptogium  myochroum  (EHRH.,  SCHAER.)  TUCK. 
Placodium  aurantiacum  (LIGHTF.)  NAEG.  and  HEPP. 
Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP. 
Lecanora  subfusca  (L.)  ACH. 
Lecanora  varia  (EHRH.)  NYL. 
Lecanora  verrucosa  (Acn.)  LAUR. 
Pertusaria  finkii  A.  ZAHLB. 
Biatora  turgidula  (FR.)  NYL. 
Biatora  arthropurpurea  (MASS.)  HEPP. 
Biatora  rubella  (EHRH.)  RABENH. 
Biatora  fuscorubella  (HOFFM.)  TUCK. 
Biatora  atrogrisea  (DELIS.)  HEPP. 
Lecidea  enteroleuca  FR. 
Buellia  alboatra  (HOFFM.)  TH.  FR. 
Buellia  parasema  (Acn.)  TH.  FR. 
Opegrapha  sp. 
Graphis  scripta  (L.)  ACH. 
Graphis  scripta  (L.)  ACH.  var.  limitata  ACH. 
Graphis  scripta  (L.)  ACH.  var.  recta  (HuMB.)  NYL. 
Arthonia  lecideella  NYL. 
Calicium  lucidum  (Tn.  FR.)  FINK. 
Coniocybe  pallida  (PERS.)  FR. 
Melaspilea  arthonioides  (FEE)  NYL. 
Pyrenula  leucoplaca  (WAHL.)  KBR. 

Pyrenula  leucoplaca  (WAHL.)  KBR.  var.  pluriloculata  var.  nov. 
I  shall  now  record  the  smooth  bark  formation  and  then  con- 
sider the  adaptations  of  the  two  groups  together. 

PYRENULA  FORMATION  OF  TREES  WITH  SMOOTH  BARK 

(BEMIDJI). 

Theloschistes  polycarpus  (EHRH.)  TUCK. 
Theloschistes  concolor  (DICKS.)  TUCK. 
Parmelia  olivacca  (L.)  ACH. 
Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP. 


672  MINNESOTA    BOTANICAL    STUDIES. 

Lecanora  subfusca  (L.)  ACH. 

Lecanora  varia  (EHRH.)  NYL. 

Rinodina  sophodes  (Acn.)  NYL. 

Biatora  atropurpurea  (MASS.)  HEPP. 

Biatora  rubella  (EHRH.)  RABENH. 

Lecidea  enteroleuca  FR. 

Buellia  parasema  (Acn.)  TH.  FR. 

Graphis  scripta  (L.)  ACH. 

Arthonia  lecideella  NYL. 

Arthonia  radiata  (PERS.)  TH.  FR. 

Sagedia  oxyspora  (NYL.)  TUCK. 

Pyrenula  punctiformis  (Acn.)  NAEG.  var.  follax  NYL. 

Pyrenula  cinerella  (FLOT.)  TUCK.  var.  quadriloculata  FINK. 

Pyrenula  leucoplaca  (WAHL.)  KBR. 

Pyrenula  leucoplaca  (WAHL.)  KBR.    var.  pluriloculata   var. 
nov. 

Inspection  of  the  list  of  plants  given  above  for  the  rough 
bark  formations  based  on  characters  of  substratum,  shows 
lichens  varying  widely  structurally  and  adapted  to  the  forma- 
tion in  very  different  ways.  There  are  the  Biatoras,  which 
usually  occur  in  the  more  moist  and  shaded  spots  in  this 
scattered  formation  and  have  small  thalli  usually  without  cellu- 
lar cortex.  The  same  may  be  said  of  the  Buellias  and  the 
Acolium,  while  the  Lecanoras,  Placodzums,  and  Pertusarias 
usually  have  somewhat  larger  thalli  and  some  indication  of  an 
upper  cortex  at  least.  Even  more  rudimentary  than  any  of  the 
above  are  the  thalli  of  the  Opegraphas,  Graphics,  Pyrenulas 
and  the  Coniocybe  which  are  hypophloeodal  and  thus  protected. 
However  the  Collemas  without  cellular  cortex  rise  to  the  foli- 
aceous  type,  but  seek  very  damp  and  well  shaded  habitats. 
The  Leptogium^  with  upper  cortex  only,  seeks  damp  places 
also.  Next  may  be  considered  the  Theloschistes,  Parmelias 
and  Physcias  with  foliaceous  thalli  having  good  cortex  on  all 
sides,  and  which  are  not  so  closely  adnate  to  the  substratum  as 
the  crustaceous  forms.  Finally  we  have  the  fruticulose  type  of 
structure  represented  in  the  formation  in  the  Ramalinas  and 
Usneas.  These  plants  have  a  good  cortex  on  all  sides  though 
not  cellular  at  least  in  the  first  genus,  and  in  well  shaded  and 
moist  portions  of  the  formation  are  quite  common.  This  for- 
mation furnishes  the  most  variety  as  to  methods  of  adaptation 
of  any  herein  recorded. 


Fink:   LICHENS  OF  NORTHWESTERN  MINNESOTA.  673 

The  lichens  composing  the  smooth  bark  formation  are  not  so 
various  in  type,  including  only  the  more  crustaceous  and  rudi- 
mentary types  of  those  given  above.  The  more  developed 
forms  with  more  specialized  rhizoids  usually  fail  to  gain  a  foot- 
hold on  the  smooth  bark. 

Next  in  order  may  be  considered  the  formation  of  old  wood. 

CALICEI    LICHEN    FORMATION    OF    DEAD    WOOD    (BEMIDJl). 

Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP.  var.  pyracea 
NYL.,  C. 

Placodium  ferrugineum  (Huos.)  HEPP. 

Lecanora  varia  (EHRH.)  NYL.,  C. 

Lecanora  varia  (EHRH.)  NYL.  var.  ssepincola  FR. 

Lecanora  varia  (EHRH.)  NYL.  var.  symmicta  ACH. 

Rinodina  sophodes  (Acn.)  NYL.,  C. 

Biatora  uliginosa  (SCHRAD.)  FR. 

Lecidea  enteroleuca  FR.  var.  ambigua  ANZ. 

Buellia  parasema  (Acn.)  TH.  FR.,  C. 

Buellia  myriocarpa  (DC.)  MUDD. 

Acolium  tigillare  (Acn.)  DN. 

Calicium  lucidum  (Tn.  FR.)  FINK. 

Calicium  sp. 

Calicium  trabinellum  (SCHAER.)  KBR. 

Calicium  parietinum  ACH. 

Calicium  trichiale  ACH.  var.  cinereum  NYL. 

This  formation  is  given  a  slightly  different  designation  from 
the  similar  ones  previously  recorded  for  Mankota  and  Granite 
Falls  *  because  of  the  absence  of  old  boards  in  the  new  country 
about  Bemidji.  The  formation  recorded  in  this  paper  is  similar 
to  the  two  formerly  recorded,  but  much  richer,  especially  in 
Calicei,  for  which  group  the  formation  is  named.  Yet  the 
three  Calicei  formations  recorded  for  the  state  show  more  varia- 
tion than  other  related  formations  and  have  only  four  (C)  com- 
mon species  of  a  total  of  23.  Moreover,  not  a  single  member 
of  the  Calicei  is  common  to  the  three  formations.  From  some 
hasty  observations  made  in  1897  in  the  Lake  Superior  region, \ 
I  had  come  to  believe  that  certain  Caliciums  normally  grow  on 
rotting  wood.  Careful  observation  in  1900  showed  that  all 

*Fink,  B.     1.  c.,  305-306. 
fFink,  B.     1.  c.,  306. 


674  MINNESOTA    BOTANICAL    STUDIES. 

found  during  that  summer  at  least  belong  to  living  or  to  dead 
wood  formations  though  occasionally  the  plants  on  dead  wood 
may  persist  after  the  wood  has  begun  to  decay  perceptibly  at 
the  surface. 

The  lichens  of  this  formation  are  of  the  small,  closely  adnate 
or  hypophoeodal  crustaceous  type  with  upper  cortical  protection 
in  the  Lecanoras  and  Placodiums  and  none  in  the  Rinodina^ 
the  Lecidea,  the  Buellias  and  the  Caliciums.  The  substratum 
is  a  dry  one  usually,  as  I  have  not  concluded  the  more  damp  and 
more  frequently  rotting  logs  of  the  tamarack  swamps,  which 
have  their  own  peculiar  formation  recorded  below.  Hence,  the 
crustaceous  type  of  thallus  is  especially  adapted  to  the  forma- 
tion. True  the  Caliciums  put  up  the  minute  erect  podetia,  but 
these  are  solid  cylinders  of  hyphae  running  in  a  longitudinal 
direction  and  usually  quite  devoid  of  the  algal  symbionts  which 
might  suffer  from  dryness. 

The  last  of  the  formations  to  be  compared  with  those  studied 
in  1897  in  southwestern  Minnesota  is  the  following. 

CLADONIA     LICHEN     FORMATION     OF    ROTTEN     WOOD     (BEMIDJl). 

Peltigera  canina  (L.)  HOFFM.,  C. 

Peltigera  canina  (L.)  HOFFM.  var.  sorediata  SCHAER. 

Cladonia  mitrula  TUCK. 

Cladonia  pyxidata  (L.)  FR. 

Cladonia  pyxidata  (L.)  FR.  var.  neglecta  (FLK.)  MASS. 

Cladonia  fimbriata  (L.)  FR.  var.  tubaeformis  FR.,  C. 

Cladonia  fimbriata  (L.)  FR.  var.  simplex  (WEIS.)  FLOT. 

Cladonia  gracilis  (L.)  NYL.,  C. 

Cladonia  gracilis  (L.)  NYL.  var.  hybrida  SCHAER. 

Cladonia  gracilis  (L.)  NYL.  var.  symphycarpia  TUCK. 

Cladonia  gracilis  (L.)  NYL.  var.  verticillata  FR.,  C. 

Cladonia  gracilis  (L.)  NYL.  var.  anthocephala  FLK. 

Cladonia  macilenta  (EHRH.)  HOFFM. 

Cladonia  bacillaris  NYL. 

Cladonia  cristatella  TUCK. 

Comparing  this  formation  with  the  corresponding  ones  at 
Mankato  and  Granite  Falls,*  we  find  that  the  formations  are 
constant  at  least  as  regards  genera,  the  one  herein  recorded  for 
a  region,  especially  rich  in  Cladonias,  being  of  course  richer  in 
species  than  the  two  formerly  studied.  The  addition  of  a  third 

*Fink,  B.     1.  c.,  306-307. 


Fink:    LICHENS  OF  NORTHWESTERN  MINNESOTA.  675 

locality  has  only  diminished  the  number  of  common  species  (C) 
by  one.  Like  the  very  similar  Cladonia-Peltigera  formation 
recorded  above,  this  one  is  especially  developed  under  the 
pines.  Indeed,  it  hardly  seems  necessary  to  separate  them  in 
the  region  now  under  consideration. 

The  adaptations  of  the  plants  of  the  formation  are  similar  to 
those  fully  discussed  under  the  Cladonia-Peltigera  formation. 
Briefly  then,  the  Peltigeras,  having  only  an  upper  cellular  cor- 
tex, lie  flat  on  the  substratum,  while  the  Cladom'as,  protected 
on  all  sides  by  a  pseudo-cortex,  rise  vertically  and  become 
luxuriant  in  the  moist  and  shaded  habitat. 

Next  in  order  I  shall  consider  a  number  of  formations  some- 
what studied  in  the  Lake  Superior  region  in  1897,  but  not  yet 
recorded  for  the  state,  as  it  was  then  found  impossible  to  take 
sufficient  data  in  the  rapid  survey  of  a  region  fully  one  fourth 
of  whose  lichen  flora  the  writer  had  previously  known  only 
through  herbarium  specimens  or  not  at  all.  In  the  second  sur- 
vey of  a  somewhat  similar  region,  I  was  able  to  take  sufficient 
field  notes  upon  which  to  base  an  ecologic  study  of  lichen  for- 
mations peculiar  to  the  region  as  well  as  those  previously 
known.  These  formations,  new  to  the  state,  will  be  recorded 
for  more  than  one  place  as  far  as  possible. 

First  of  all  I  shall  record  the  formation  of  the  pine  trees, 
naming  it  for  the  Usnei  group  which  give  character  to  this 
plant  community : 

USNEI    LICHEN    FORMATION    OF    THE    PINES  (RED    LAKE). 

Cetraria  ciliaris  (Acn.)  TUCK,  (cones). 

Cetraria  juniperina  (L.)  ACH.  var.  pinastri  ACH. 

Evernia  prunastri  (L.)  ACH.  (cones). 

Usnea  barbata  (L.)  FR.  var.  florida  Fr. 

Alectoria  jubata  (L.)  TUCK.  var.  chalybeiformis  ACH. 

Theloschistes  chrysopthalmus  (L.)  NORM. 

Parmelia  physodes  (L.)  ACH. 

Physcia  tribacia  (Acn.)  TUCK. 

Physcia  hispida  (SCHREB.,  FR.)  TUCK. 

Lecanora  varia  (EHRH.)  NYL.  (cones). 

Lecanora  subfusca  (L.)  ACH.  var.  argentata  (Acn.)  (cones). 

Buellia  parasema  (Acn.)  TH.  FR.  (cones). 

Calicium  lucidum  (TH.  FR.)  FINK. 

On  approaching  a  pine  woods  one  is  impressed  with  an  ap- 


676  MINNESOTA    BOTANICAL    STUDIES. 

parent  dearth  of  lichens,  but  after  a  careful  study  he  becomes 
convinced  that  the  plants  are  common  enough  when  the  trees, 
the  fallen  branches  and  the  earth  are  carefully  observed.  The 
foliaceous  Parmelias,  Physcias,  etc.,  so  common  on  the  decidu- 
ous trees,  are  not  so  frequently  seen,  but  the  pines  certainly 
have  their  own  peculiar  formations,  composed  of  lichens  found 
of  course  on  other  substrata  in  various  parts  of  the  state.  The 
variety  of  Cetraria  juniperina  (L.)  Ach.  and  the  Calicium  lu- 
cilium  (Th.  Fr.)  Fink  are  perhaps  the  most  characteristic  ele- 
ments of  the  formation,  as  they  are  seldom  seen  elsewhere  than 
on  the  conifers,  though  by  no  means  common  even  in  this  for- 
mation. The  similar  formation  was  studied  at  Bemidji,  and 
the  only  difference  is  that  Lecanora  subfusca  (L.)  Ach.  replaces 
the  variety.  As  indicated  in  the  list  of  species,  a  number  of  the 
plants  are  as  common  or  more  so  on  the  old  cones  as  on  other 
portions  of  the  trees.  Careful  comparison  of  this  formation 
with  the  Parmelei  formation  of  trees  with  rough  bark  will  dem- 
onstrate that  the  two  are  quite  distinct,  though  resembling  each 
other  in  some  respects. 

The  adaptations  of  the  lichens  of  this  formation  are  some- 
what various.  They  have  been  discussed  somewhat  under  the 
rough  bark  formation  above  and  will  be  further  noticed  under 
the  Usnea  formation  below.  However,  while  it  is  apparent  that 
the  fruticulose  lichens  of  the  Usnei  group  seem  well  adapted  to 
the  swamp  Usnea  formation  given  below  where  moisture  is 
abundant,  I  have  not  been  able  to  satisfy  myself  that  the  pines 
of  the  higher  ground  furnish  more  moisture  than  the  deciduous 
trees  of  similar  grounds.  I  suspect  that  the  plants  in  this  in- 
stance and  in  the  formation  given  below  for  the  swamps  are 
quite  as  much  influenced  in  their  choice  of  habitat  by  an  adap- 
tation to  coniferous  wood  as  by  amount  of  moisture.  And  I 
may  add  here  that  in  other  portions  of  this  paper  I  have  not 
taken  into  account  any  adaptations  of  the  lichens  to  substrata  of 
certain  physical  or  chemical  composition.  The  subject  is  a 
difficult  one  as  yet  little  understood,  and  moreover  I  am  con- 
vinced from  observation  that,  beyond  the  well-known  fact  that 
some  lichens  prefer  rocks,  calcareous  in  some  instances  and 
granitic,  etc.,  in  others,  while  others  prefer  trees  and  in  some 
instances  a  particular  species  or  genus,  the  conditions  as  to  shade, 
moisture,  etc.,  are  the  important  ones  in  determining  the  habitats 
of  lichens  and  the  composition  of  lichen  formations. 


Fink:    LICHENS  OF  NORTHWESTERN  MINNESOTA.  677 

I  studied  the  lichens  of  the  earth  and  rotting  wood  under  the 
pines  with  a  view  to  establishing  a  formation  ;  but  the  plants 
were  found  to  be  so  similar  to  those  of  the  Cladonia  lichen  for- 
mation of  rotten  wood  and  the  Cladonia- Pelt  iger  a  lichen  for- 
mation of  shaded  earth  previously  established  that  I  have 
recorded  them  above  in  these  formations,  though  possibly  I 
have  done  violence  in  not  separating  them  in  this  region  for  the 
sake  of  correlating  results  with  former  work  in  another  area. 

Because  of  close  relationship  to  the  above  formation,  I  shall 
next  consider  the  Usnea  formation. 

USNEA  LICHEN  FORMATION  OF  TAMARACKS  IN  SWAMPS 
(HENNING). 

Cetraria  ciliaris  (Acn.)  TUCK.,  C. 

Evernia  prunastri  (L.)  ACH.,  C. 

Usnea  barbata  (L.)  FR.  var.  florida  FR.,  C. 

Usnea  barbata  (L.)  FR.  var.  ceratina  SCHAER. 

Usnea  barbata  (L.)  FR.  var.  dasypoga  FR. 

Usnea  cavernosa  TUCK.,  C. 

Alectoria  jubata  (L.)  TUCK.  var.  chalybeiformis  ACH.,  C. 

Parmelia  physodes  (L.)  ACH.,  C. 

Parmelia  olivacea  (L.)  ACH.,  C. 

Parmelia  caperata  (L.)  ACH.,  C. 

Physcia  hispida  (Acn.)  TUCK.,  C. 

Calicium  trichiale  ACH.  var.  cinereum  NYL. 

The  formation  was  afterward  carefully  examined  at  Bemidji 
and  Red  lake  and  was  found  to  be  a  very  characteristic  one. 
At  Bemidji  were  found  all  of  the  above-marked  (C)  and  Rama- 
lina  pusilla  (Prev.)  Tuck.,  Cetraria  juniperina  (L.)  Ach.  var. 
pinastri  Ach.,  Alectoria  jubata  (L.)  Tuck,  and  Calicium 
chrysocephalum  Ach.  in  addition.  The  Red  lake  formation 
contained  all  of  the  species  found  at  Bemidji  except  the  last  two, 
so  that  those  marked  (C)  are  the  ones  common  to  the  three 
similar  formations. 

The  dominant  elements  in  this  formation  are  the  Usneas,  the 
Evernia  and  the  Alectoria,  which  in  the  moist  swamps  and  pio- 
tected  on  all  sides  by  a  pseudo-cortex  of  closely  arrar  ged 
hyphae,  hang  suspended  or  grow  up  from  the  substrati  ,n  in 
fruticulose  fashion.  With  these  occur  the  less  charac  eristic 
elements  whose  structure  has  been  fully  discussed  ar  .  whose 
adaptations  may  be  easily  inferred.  Though,  as  st?'  d  in  the 


678  MINNESOTA    BOTANICAL    STUDIES. 

discussion  of  the  Usnei  formation  of  the  pines,  the  plants  may 
be  adapted  to  the  coniferous  wood  as  well  as  to  the  moist  habitat, 
it  remains  to  be  stated  that  the  same  species  are  both  more 
numerous  and  more  luxuriant  in  the  swamps  than  on  the  upland 
pines. 

The  following  formation  must  be  regarded  as  somewhat 
doubtful  till  further  studied. 

STICTA  PULMONARIA   LICHEN    FORMATION    OF    CEDAR  SWAMPS 

(BEMIDJI). 

Sticta  pulmonaria  (L.)  ACH. 

Nephroma  leevigatum  ACH. 

Lecanora  pallida  (SCHREB.)  SCHAER. 

Lecanora  pallescens  (L.)  SCHAER. 

Pertusaria  communis  DC. 

Calicium  chrysocephalum  ACH. 

There  is  no  doubt  of  the  distinctness  of  the  formation  in 
the  region  as  none  of  the  plants  were  found  elsewhere  during 
the  summer  than  in  the  cedar  swamps,  except  the  last  which 
was  also  found  in  the  tamarack  swamps.  The  plants  of  the 
formation  grow  in  dryer  places  in  other  regions  and  seem  to  be 
as  well  adapted  to  upland  woods  as  many  other  lichens  recorded 
for  the  tree  formations  of  higher  ground. 

The  swamps  afford  yet  one  other  characteristic  formation 
well  developed  in  the  area  studied  wherever  tamaracks  and 
cedars  flourish.  It  may  appropriately  be  designated  as  follows:' 

THE  CALICIUM  LICHEN  FORMATION  OF  OLD  LOGS  AND  STUMPS  IN 
TAMARACK    SWAMPS  (HENNING). 

Ramalina  pusilla  (PREV.)  TUCK.,  C. 

Cetraria  ciliaris  (Acn.)  TUCK.,  C. 

Parmelia  saxatilis  (L.)  FR.,  C. 

Cladonia  mitrula  TUCK.,  C. 

Cladonia  fimbriata  (L.)  FR.  var.  tubaeformis  FR.,  C. 

Cladonia  gracilis  (L.)  NYL.,  C. 

Cladonia  gracilis  (L.)  NYL.  var.  hybrida  SCHAER.,  C. 

Calicium  trichiale  ACH.  var.  cinereum  NYL.,  C. 

Calicium  trachelinum  ACH. 

Calicium  sp. 

Calicium  curtum  TURN,  and  BORR. 


Fink:   LICHENS  OF  NORTHWESTERN  MINNESOTA.  679 

Calicium  trabinellum  (SCHAER.)  KBR. 

Calicium  parietinum  ACH. 

The  similar  formations  were  studied  at  Bemidji  and  Red 
lake,  and  the  species  marked  (C)  were  found  in  each  of  the 
three  formations.  For  Bemidji  may  be  added  Biatora  virides- 
cens  (Schrad.)  Fr,  and  for  both  Bemidji  and  Red  Lake,  Cetraria 
juniperina  (L.)  Ach.  var.  pinastri  Ach.  Although  occurring 
in  the  same  area  as  the  second  formation  above  on  the  living 
tamaracks,  the  present  one  will  be  found  by  comparison  to  be 
quite  distinct  both  as  to  genera  and  species. 

Growing  in  moist  areas,  the  lichens  of  the  formation  are 
almost  uniformly  those  which  rise  more  or  less  from  the  sub- 
stratum. Nearly  all  are  the  Cladonias  and  Caliciums,  which 
have  podetia  rising  erect  from  the  substratum  and  are  protected 
on  all  sides  by  a  pseudo-cortex  of  densely  interwoven  hyphae 
running  in  a  longitudinal  direction. 

The  earth  lichen  formations  of  the  swamps  were  carefully 
noted  and  were  found  to  be  essentially  like  those  of  Cladonia 
formations  of  rotten  wood  recorded  above,  at  least  as  to  genera. 
I  shall  record  the  formation  provisionally  that  it  may  be  further 
studied  and  shall  name  it  for  a  variety  of  Peltigera  thus  far 
found  in  the  state  only  in  the  swamps. 

PELTIGERA    CANINA    LEUCORRHIZA    LICHEN    FORMATION    OF 
EARTH    IN    TAMARACK    SWAMPS    (BEMIDJl). 

Peltigera  canina  (L.)  HOFFM.,  C. 

Peltigera  canina  (L.)  HOFFM.  var.  leucorrhiza  FLK.,  C. 

Cladonia  cariosa  (AcH.)  SPRENG.,  R  L. 

Cladonia  pyxidata  (L.)  FR.,  C. 

Cladonia  gracilis  (L.)  NYL.  var.  verticillata  FR.,  C. 

Cladonia  gracilis  (L.)  NYL.  var.  hybrida  SCHAER.,  C. 

Cladonia  cenotea  (Acn.)  SCHAER.,  R  L. 

Cladonia  furcata  (Huos.)  FR.,  B  and  H. 

Cladonia  rangiferina  (L.)  HOFFM.,  C. 

The  similar  formations  were  studied  at  Henning  and  Red 
lake.  Those  marked  (C)  are  common  to  the  three  formations, 
those  marked  (R  L)  were  found  at  Red  lake  only  and  the  one 
marked  (B  and  H)  at  Henning  as  well  as  at  Bemidji.  The 
adaptations  of  the  plants  of  these  formations  are,  of  course,  the 
same  as  those  of  the  lichens  of  the  Cladonia  formations  of 
rotten  wood. 


680  MINNESOTA    BOTANICAL    STUDIES. 

The  following  formation  was  distinctly  discernible  in  the  low 
woods  about  the  tamarack  swamps  near  Henning,  the  species 
composing  it  being  unusually  abundant.  From  the  frequence 
of  Pertusarias  rare  elsewhere  in  the  region  studied  during  the 
summer,  I  shall  name  it  as  follows  : 

PERTUSARIA  LICHEN  FORMATION  OF  TREES  IN  LOW  WOODS 

(HENNING). 

Physcia  obscura  (EHRH.)  NYL. 

Collema  flaccidum  ACH. 

Collema  nigrescens  (HUBS.)  ACH. 

Leptogium  myochroum  (EHRH.,  SCHAER.)  TUCK. 

Pertusaria  velata  (TURN.)  NYL. 

Pertusaria  pustulata  (Acn.)  NYL. 

Biatora  glauconigrans  TUCK. 

Biatora  rubella  (EHRH.)  RAB. 

Biatora  varians  (Acn.)  TUCK. 

I  failed  to  find  such  a  formation  elsewhere  in  the  territory 
explored.  The  plants  in  the  formation  grow  on  the  common 
deciduous  trees  of  the  area,  whereas  at  Bemidji  and  Red  lake 
the  swamps  were  surrounded  for  the  most  part  by  pines.  As  a 
whole  the  various  adaptations  of  the  plants  are  not  difficult  to 
detect.  It  is  a  little  peculiar  that  the  species  of  Physcia,  be- 
longing to  this  damp  formation,  belongs  to  the  section  of  the 
genus  having  a  well  developed  parenchymatous  cortex  rather 
than  to  the  one  having  a  non-cellular  cortex  of  closely  packed 
hyphae.  The  Cottemas  with  their  non-cortical  thallus  and  the 
Leptogium  with  a  cortex  of  a  single  layer  of  cells  are,  of  course, 
quite  at  home  in  such  a  damp  habitat,  as  are  the  Biatoras  with 
their  thallus  devoid  of  cortex.  The  Pertusarias  have  a  fairly 
developed  upper  cortex. 

The  last  formation  to  be  recorded  is  a  scattered  one  detected 
in  shaded  places  at  Bemidji,  which  may  be  designated  as 
follows  : 

BIATORA    LICHEN    FORMATION    OF    MOSSES    (fiEMIDJl). 

Pannaria  languinosa  (Acn.)  KBR. 
Biatora  vernalis  (L).  Fr. 
Biatora  sphaeroides  (DICKS.)  TUCK. 
Biatora  hypnophila  (TURN.)  TUCK. 


Fink:    LICHENS    OF    NORTHWESTERN    MINNESOTA.  681 

The  first  and  last  Biatoras  are  the  common  elements  of  the 
formation  and  may  be  found  in  this  habitat  commonly  at  Be- 
midji.  The  last  Biatora  is  the  only  plant  of  the  formation 
noticed  elsewhere  in  such  environment,  nor  were  the  other  two 
Biatoras  found  elsewhere  during  the  summer.  The  formation 
was  observed  in  the  Lake  Superior  region  at  Gunflint,  where 
the  second  Biatora  was  wanting,  and  in  the  Snowbank  lake 
area,  where  the  first  one  did  not  occur.  The  plants  of  the  for- 
mation all  have  rudimentary  thalli  devoid  of  cortical  layers  and 
are  well  adapted  to  the  moist  shaded  habitat  on  the  damp  shaded 
sides  of  tree  bases  where  they  spread  over  the  mosses. 

Nearly  all  of  the  formations  herein  recorded  are  more  or  less 
scattered  in  the  sense  explained  in  the  fifth  paper  of  this  series.* 
In  this  paper  as  in  the  others  I  have  avoided  attempting  too 
close  analysis  as  to  amount  of  illumination,  roughness  of  bark, 
amount  of  moisture,  and  have  omitted  from  the  lists  of  plants  of 
the  various  formations  those  rarer  lichens  whose  adaptations 
seemed  most  doubtful.  Yet  in  attempting  a  detailed  study  of  a 
single  group  of  plants,  I  feel  sure  that,  if  I  have  erred  at  all,  it 
has  been  in  including  some  doubtful  elements  in  a  few  of  the 
formations.  In  general,  I  have  found  that  one  can  attempt  an 
amount  of  minute  detail  in  such  a  study  which  could  only  be 
carried  out  by  a  long  study  of  a  single  locality  and  which  would 
probably  not  be  more  helpful  than  such  general  survey  as  I  feel 
that  I  have  been  able  to  conduct  in  the  field  with  some  degree 
of  success. 

Notwithstanding  the  recording  of  16  distinct  formations  for 
the  region  now  under  consideration  and  only  14  for  southwest- 
ern Minnesota,  I  still  adhere  to  the  statement  already  recorded 
in  this  paper  that  the  conditions  of  lichen  growth  are  more  uni- 
form in  the  former  area.  The  greater  diversity  in  the  latter  ter- 
ritory may  be  seen  in  the  fact  that  there  is  a  larger  amount  of 
difference  between  similar  formations  in  the  various  parts  of  the 
region,  due  to  more  variation  in  amount  of  moisture,  shade, 
etc.,  and  in  the  circumstance  that  some  of  the  formations  bear- 
ing different  names  in  the  former  region  are  very  much  alike. 
On  account  of  this  greater  uniformity  of  conditions  under  which 
similar  formations  seems  to  have  developed,  it  has  been  even 
more  difficult  than  in  the  preceding  papers  to  ascertain  why  cer- 
tain species  are  found  in  a  formation  at  one  place  and  not  in  the 

*Fink,  B.     1.  c.,  307. 


682  MINNESOTA    BOTANICAL    STUDIES. 

similar  formation  at  another.  I  have,  consequently,  seldom  at- 
tempted such  explanations  in  this  paper. 

In  the  present  paper,  after  a  more  extended  study  of  lichen 
formations  in  the  field,  I  have  attempted  a  more  minute  discus- 
sion of  structural  adaptations  based  upon  careful  study  of  thalli 
in  the  laboratory.  This  analysis  has  not  in  most  instances  de- 
tracted from  the  apparent  genuineness  of  the  formations,  though 
in  some  it  has  not  been  possible  to  show  that  every  plant  is 
structurally  adapted  to  the  formation  in  which  it  occurs. 

Throughout  this  paper  I  have  referred  only  once  to  the  in- 
fluence of  physical  structure  and  chemical  composition  of  sub- 
strata as  influencing  the  distribution  of  lichens  and  the  compo- 
sition of  lichen  formations.  In  the  beginning,  doubtless,  lichen 
species  were  influenced  in  their  choice  of  substrata  by  their 
adaptations  to  light,  shade,  moisture  and  other  conditions, 
though  of  course  such  physical  conditions  of  substrata  as  in- 
fluence transfer  of  moisture  are  also  to  be  considered  as  they 
have  been  in  my  discussions,  as  have  also  roughness  and 
smoothness  of  surface.  No  doubt  both  physiological  and  an- 
atomical changes  frequently  result  in  lichens  from  adoption  of 
certain  substrata,  but  it  is  well  known  that  the  conditions  of 
life  in  lichens  are  such  that  they  are  not  so  much  dependent 
upon  or  influenced  by  their  substrata  as  are  most  other  plants. 
Thus  a  large  proportion  of  lichens  occur  commonly  upon  sub- 
strata of  the  most  varied  chemical  composition  and  physical 
structure,  provided  the  conditions  of  light,  moisture,  etc.,  named 
above  are  favorable.  Therefore  these  factors  must,  for  the 
present  at  least,  receive  chief  attention  in  the  study  of  ecologic 
distribution  of  lichens,  though  the  more  difficult  and  less  im- 
portant subject  of  the  influence  of  physical  and  chemical  make 
up  of  substrata  is  well  worth  attention.  Doubtless  in  the 
struggle  of  lichen  species  for  possession  of  substrata,  adapta- 
tions to  chemical  composition  of  substrata  sometimes  play  an 
important  part.  For  instances,  the  crustaceous  lichens  of  the 
calcareous  rocks  and  earth  produce  a  fat  which  is  probably 
utilized  by  the  plants  for  purposes  of  nutrition.  Hence,  as 
these  plants  can  build  up  the  fats  from  material  obtained  wholly 
or  in  part  from  the  calcareous  substrata,  they  would  have  an 
advantage  over  lichens  which  can  not  thus  utilize  the  carbon 
of  the  rocks,  in  the  struggle  for  possession.  Lichens  produce 
other  chemical  compounds,  some  of  which  are  doubtless  depen- 


Fink  :     LICHENS    OF    NORTHWESTERN    MINNESOTA.  683 

dent  upon  the  nature  of  the  substratum.  To  what  extent  these 
compounds  are  of  use  to  the  plant,  or  in  what  degree  they  are 
derived  from  the  substratum  is  little  known.  Till  these  prob- 
lems are  solved  we  can  hardly  hope  to  discuss  intelligently  the 
influence  of  chemical  composition  upon  distribution. 

I  am  under  obligations  to  Dr.  A.  Zahlbruckner,  of  Vienna, 
Austria,  for  aid  in  the  determination  of  several  of  the  species 
listed  below,  and  also  to  Dr.  E.  Wainio,  of  Helsingfors,  Fin- 
land, for  examination  and  determination  of  the  larger  part  of 
the  Cladonias. 

LIST    OF    SPECIES    AND    VARIETIES. 

1.  Ramalina  calicaris  (L.)  FR.  var.  fraxinea  FR. 

On  trees,  infrequent.  Battle  lake,  June  20,  1900,  no.  39. 
Henning,  June  28,  1900,  no.  328  and  July  2,  1900,  no.  412. 
Thief  River  Falls,  July  18,  1900,  no.  796  and  July  23,  1900, 
no.  869.  Red  lake,  July  26,  1900,  no.  882  and  July  31, 
1900,  no.  1018. 

2.  Ramalina  calicaris  (L.)  FR.  var.  fastigiata  FR. 

On  trees,  frequent.  Battle  lake,  June  18,  1900,  no.  2. 
Leaf  hills,  June  26,  1900,  no.  209  and  June  27,  1900,  no.  246. 
Henning,  June  28,  1900,  no.  330  and  332  and  July  2,  1900,  no. 
411.  Bemidji,  July  5,  1900,  no.  471.  Thief  River  Falls,  July 
17,  1900,  no.  762.  Red  lake,  July  26,  1900,  no.  879  and 
July  31,  1900,  no.  1006. 

3.  Ramalina  pusilla  (PREV.)  TUCK. 

On  tamarack  in  swamps,  rare.  Henning,  June  30,  1900,  no. 
370.  Bemidji,  July  7,  1900,  no.  517  and  July  8,  1900,  no. 
531.  Red  lake,  July  31,  1900,  no.  1012. 

4.  Cetraria  ciliaris  (AcH.)  TUCK. 

Common  on  pines  and  on  tamaracks  in  swamps.  Battle 
lake,  June  20,  1900,  no.  71.  Henning,  June  25,  1900,  no. 
188,  June  28,  1900,  nos.  315  and  323  and  June  30,  1900,  no. 
378.  Bemidji,  July  4,  1900,  no.  427,  July  5,  1900,  nos.  472, 
476  and  490,  July  7,  1900,  nos.  522  and  526,  July  12,  1900, 
no.  680  and  July  14,  1900,  no.  730.  Red  lake,  July  28,  1900, 
no.  463,  July  30,  1900,  nos.  985  and  988,  August  2,  1900,  no. 
1050  and  August  3,  1900,  no.  1070. 

5.  Cetraria  juniperina  (L.)  ACH.  var.  pinastri  ACH. 

On  pines  and  on  tamaracks  in  swamps,  rare.     Bemidji,  July 


684  MINNESOTA    BOTANICAL    STUDIES. 

7,  1900,  no.  516,  July  9,  1900,  no.  543  and  July  12,  1900,  no. 
665.  Red  lake,  July  27,  1900,  no.  925  and  August  4,  1900, 
no.  1083. 

6.  Evernia  prunastri  (L.)  ACH. 

Common  on  pines  and  abundant  on  tamaracks  in  swamps. 
Battle  lake,  June  23,  1900,  nov  140.  Henning,  June  25,  1900, 
no.  187  and  June  28,  1900,  no.  312.  Bemidji,  July  4,  no.  428, 
July  6,  1900,  no.  518  and  July  7,  1900,  no.  531.  Thief  River 
Falls,  July  19,  1900,  no.  827.  Red  lake,  July  28,  1900,  no. 
970. 

7.  Usneabarbata  (L.)  FR. 

On  trees,  frequent.      Henning,  June  28,  1900,  no.  513. 

8.  Usnea  barbata  (L.)  FR.  var.  florida  FR. 

On  trees,  frequent.  Battle  lake,  June  19,  1900,  no.  27. 
Henning,  June  25,  1900,  no.  174.  Leaf  hills,  June  27,  1900, 
no.  248.  Bemidji,  July  4,  1900,  no.  425,  July  7,  1900,  no. 
532  and  July  12,  1900,  no.  666.  Thief  River  Falls,  July  20, 
1900,  no.  838.  Red  lake,  July  28,  1900,  no.  945  and  August 
2,  1900,  no.  1053. 

9.  Usnea  barbata  (L.)  FR.  var.  ceratina  SCHAER. 

On  trees,  frequent.     Henning,  June  30,    1900,  no.  375. 

10.  Usnea  barbata  (L.)  FR.  var.  dasypoga  FR. 

On  trees,  abundant.      Henning,  June  25,  1900,  no.  191. 

u.  Usnea  cavernosa  TUCK. 

On  trees,  frequent.  Henning,  June  25,  1900,  nos.  182,  192 
and  193,  June  28,  1900,  nos.  298  and  337  and  July  2,  1900,  no. 
410.  Bemidji,  July  5,  1900,  nos.  469,  470  and  488,  July  7, 
1900,  nos.  505,  513  and  519,  July  8,  1900,  nos.  534,  536  and 
544,  July  9,  1900,  no.  546  and  July  13,  1900,  no.  715.  Red 
lake,  July  27,  1900,  nos.  910,  912  and  920,  July  28,  1900,  no. 
942  and  July  30,  1900,  nos.  978,  987  and  990. 

11.  Alectoria  jubata  (L.)  TUCK. 

On  cedars,  frequent.  Bemidji,  July  6,  1900,  nos.  515  and 
520  and  July  13,  1900,  no.  701. 

13.  Alectoria  jubata  (L.)  TUCK.  var.  chalybeiformis  ACH. 

On  pines  and  tamaracks,  infrequent.  Henning,  June  28, 
1900,  no.  338.  Bemidji,  July  4,  1900,  no.  426,  July  5,  1900, 
no.  473,  July  7,  1900,  no.  512,  July  8,  1900,  no.  543  and  July 


Fink:   LICHENS  OF  NORTHWESTERN  MINNESOTA.  685 

12,  1900,  no.  683.     Red  lake,  July  28,  1900,  no.  959  and  July 
30,  1900,  no.  999. 

14.  Theloschistes  chrysophthalmus  (L.)  NORM. 

On  trees,  rare.  Battle  lake,  June  18,  1900,  no.  6.  Bemidji, 
July  ii,  1900,  no.  619.  Red  lake,  August  i,  1900,  no.  1047. 

15.  Theloschistes  polycarpus  (EHRH.)  TUCK. 

On  trees,  frequent.  Battle  lake,  June  19,  1900,  no.  35  and 
June  20,  1900,  no.  40.  Leaf  hills,  June  26,  1900,  no.  206. 
Bemidji,  July  5,  1900,  nos.  468  and  489,  July  n,  1900,  no. 
653  and  July  12,  1900,  no.  675.  Thief  River  Falls,  July  18, 
1900,  no.  786.  Red  lake,  July  27,  1900,  no.  926. 

16.  Theloschistes  lychneus  (NYL.)  TUCK. 

On  trees,  abundant  and  rarely  on  rocks.  Battle  lake,  June 
19,  1900,  no.  34.  Bemidji,  July  n,  1900,  nos.  628  and  654. 
Thief  River  Falls,  July  17,  1900,  no.  773  and  July  20,  1900, 
no.  842.  Red  lake,  July  26,  1900,  no.  884,  July  28,  1900, 
no.  964  and  August  i,  1900,  no.  1045. 

17.  Theloschistes  concolor  (DICKS.)  TUCK. 

On  trees,  frequent.  Battle  lake,  June  18,  1900,  no.  i6a. 
Leaf  hills,  June  27,  1900,  no.  265.  Thief  River  Falls,  July 

19,  1900,  no.  826.     Red  lake,  July  27,  1900,  no.  930. 

18.  Theloschistes  concolor  ( DICKS.)  TUCK.  var.  effusa  TUCK. 
On  trees,  rare.     Henning,    June  25,    1900,   no.   175.      Be- 
midji, July  5,  1900,  no.  460. 

19.  Parmelia  crinita  ACH. 

On  trees,  rare.     Bemidji,  July  12,  1900,  no.  691. 

20.  Parmelia  tiliacea  (HOFFM.)  FLK. 

In  trees,  infrequent.  Battle  lake,  June  18,  1900,  no.  16. 
Leaf  hills,  June  26,  1900,  no.  208.  Bemidji,  July  4,  1900,  no. 
449.  Red  lake,  July  31,  1900,  no.  1003. 

21.  Parmelia  borreri  TURN. 

On  trees,  frequent  and  rarely  on  rocks.     Battle  lake,  June 

20,  1900,  no.  43  and  June  21,  1900,  no.  114.     Leaf  hills,  June 
27,  1900,  nos.  281  and  286.     Bemidji,  July  11,  1900,  nos.  647 
and  658.     Thief  River  Falls,  July  19,   1900,  no.  824.     Red 
lake,  July  26,  1900,  nos.  891  and  893. 

22.  Parmelia  saxatilis  (L.)  FR. 

Common  on  trees  and  rare  on  rocks.     Battle  lake,  June  21, 


686  MINNESOTA    BOTANICAL    STUDIES. 

1900,  no.  112.  Henning,  June  25,  1900,  no.  190  and  June  30, 
1900,  no.  379.  Bemidji,  July  4,  1900,  no.  447,  July  6,  1900, 
no.  514,  July  7,  1900,  no.  521,  July  10,  1900,  no.  583,  July  n, 
1900,  nos.  646  and  660.  Thief  River  Falls,  July  19,  1900,  no. 
820  and  July  23,  1900,  no.  874.  Red  lake,  July  26,  1900, 
no.  89ia  and  August  i,  1900,  no.  1038. 

23.  Parmelia  saxatilis  (L.)  FR.  var.  sulcata  NYL. 

On  trees  and  rocks,  and  more  frequently  on  old  logs,  fre- 
quent. Battle  lake,  June  22,  1900,  no.  134.  Bemidji,  July 
4,  1900,  no.  525  and  July  7,  1900,  no.  525.  Thief  River  Falls, 
July  19,  1900,  no.  813.  Red  lake,  August  i,  1900,  no.  1034 
and  August  2,  1900,  no.  1049. 

24.  Parmelia  physodes  (L.)  ACH. 

On  pines  and  tamarack,  rare.  Henning,  June  28,  1900,  no. 
331.  Bemidji,  July  4,  1900,  no.  429  and  July  7,  1900,  no.  514. 
Red  lake,  July  27,  1900,  no.  913  and  August  3,  1900,  no. 
1071. 

25.  Parmelia  olivacea  (L.)  ACH. 

On  trees,  frequent,  and  rarely  on  rocks.  Battle  lake,  June 
19,  1900,  no.  31.  Leaf  hills,  June  26,  1900,  no.  243.  Bemidji, 
July  4,  1900,  nos.  443  and  446  and  July  7,  1900,  no.  513. 
Thief  River  Falls,  July  20,  1900,  no.  830.  Red  lake,  July 

26.  1900,  no.  885,  July  30,  1900,  no.  991   and  August  i,  1900, 
no.  1039. 

26.  Parmelia  olivacea  (L.)  ACH.  var.  aspidota  ACH. 

On  trees,   common   locally.      Leaf  hills,  July  2,   1900,  no. 

385. 

Not  previously  reported  from  Minnesota. 

27.  Parmelia  conspurcata  (SCHAER.)  WAINIO. 

On  trees,  locally  common.  Bemidji,  July  12,  1900,  nos. 
672,  682  and  700. 

Not  previously  reported  from  Minnesota. 

28.  Parmelia  caperata  (L.)  ACH. 

Common  on  trees  and  rare  on  rocks.  Battle  lake,  June  19, 
1900,  no.  26.  Henning,  June  25,  1900,  no.  195  and  July  2, 
1900,  no.  409.  Bemidji,  July  7,  1900,  no.  527  and  July  u, 
1900,  no.  659.  Thief  River  Falls,  July  20,  1900,  nos.  832, 
833  and  841.  Red  lake,  July  26,  1900,  no.  898  and  August 
i,  1900,  no.  1033. 


Fink :  LICHENS  OF  NORTHWESTERN  MINNESOTA.          687 

29.  Parmelia  conspersa  (EHRH.)  ACH. 

On  granitic  bowlders,  rare.  Bemidji,  July  16,  1900,  no.  745. 
Thief  River  Falls,  July  10,  1900,  no.  8323. 

30.  Physcia  speciosa  (WULF.,  ACH.)  NYL. 

On  bowlders  and  trees,  rare.  Bemidji,  July  n,  1900,  nos. 
598  and  613. 

31.  Physcia  hypoleuca  (MUHL.)  TUCK. 

On  trees,  rare.     Bemidji,  July  5,  1900,  no.  484. 

32.  Physcia  granulifera  (ACH.)  TUCK. 

On  trees,  rare.     Battle  lake,  June  21,  1900,  no.  115. 

33.  Physcia  pulverulenta  (SCHREB.)  NYL. 

On  trees  and  rocks,  frequent.  Battle  lake,  June  18,  no.  9. 
Bemidji,  July  9,  1900,  no.  556  and  July  n,  1900,  nos.  618  and 
657.  Thief  River  Falls,  July  17,  1900,  no.  777,  and  July  18, 
1900,  no.  793.  Red  lake,  July  26,  1900,  no.  900,  August  i, 
1900,  no.  1044  and  August  2,  1900,  no.  1051. 

34.  Physcia  stellaris  (L.)  TUCK. 

Abundant  on  trees  and  rare  on  rocks.  Battle  lake,  June  19, 
1900,  no.  23  and  June  20,  1900,  no.  93.  Leaf  hills,  June  26, 
1900,  no.  245.  Bemidji,  July  4,  1900,  no.  445  and  July  u, 
1900,  nos.  614  and  656.  Thief  River  Falls,  July  18,  1900,  nos. 
708  and  803  and  July  29,  1900,  nos.  836  and  859.  Red  lake, 
July  26,  1900,  no.  886. 

35.  Physcia  stellaris  (L.)  TUCK.  var.  apiola  NYL. 

On  rocks,  rare.  Leaf  hills,  June  26,  1900,  no.  212  and  Red 
lake,  August  i,  1900,  no.  1029. 

36.  Physcia  astroidea  (FR.)  NYL. 

On  rocks,  once  collected.  Red  lake,  August  i,  1900,  no. 
1032. 

Not  previously  reported  from  Minnesota. 

37.  Physcia  tribacia  (Acn.)  TUCK. 

On  pines,  rare.  Bemidji,  July  n,  1900,  nos.  627  and  661. 
Red  lake,  July  27,  1900,  no.  917. 

38.  Physcia  hispida  (SCHREB.,  FR.)  TUCK. 

On  trees  and  rocks,  rare.  Henning,  June  28,  1900,  no.  335. 
Bemidji,  July  u,  1900,  nos.  632  and  655.  Thief  River  Falls, 
July  18,  1900,  nos.  781  and  791.  Red  lake,  July  28,  1900,  no. 
972  and  August  2,  1900,  no.  1064. 


688  MINNESOTA    BOTANICAL    STUDIES. 

39.  Physcia  caesia  (HOFFM.)  NYL. 

On  rocks,  rare.  Battle  lake,  June  23,  1900,  no.  154.  Leaf 
hills,  June  26,  1900,  nos.  203  and  239.  Thief  River  Falls, 
July  18,  1900,  no.  787.  Red  lake,  August  4,  1900,  no.  1078. 

40.  Physcia  obscura  (EHRH.)  NYL. 

On  trees,  common,  and  rarely  on  rocks.  Battle  lake,  June 
18,  1900,  no.  4  and  June  21,  1900,  no.  121.  Leaf  hills,  June 
26,  1900,  no.  211  and  June  27,  1900,  no.  270.  Henning,  June 
29,  1900,  no.  347  and  June  30,  1900,  no.  380.  Bemidji,  July 
5,  1900,  no.  461,  July  n,  1900,  no.  622  and  July  12,  1900,  no. 
690.  Thief  River  Falls,  July  18,  1900,  no.  782,  July  19,  1900, 
no.  828  and  July  28,  1900,  no.  952.  Red  lake,  Aug.  i,  1900, 
nos.  1027  and  1043  and  Aug.  2,  1900,  no.  1065. 

41.  Physcia  adglutinata  (FLK.)  NYL. 

On  trees,  infrequent.     Red  Lake,  Aug.  i,  1900,  no.  1048. 

42.  Sticta  pulmonaria  (L.)  ACH. 

On  cedars  in  swamp,  rare.     Bemidji,  July  14,  1900,  no.  727. 

43.  Nephroma  laevigatum  ACH. 

On  old  cedars,  in  swamp,  rare.     Bemidji,  July  14,  1900,  no. 

723- 

44.  Peltigera  horizontalis  (L.)  HOFFM. 

On  earth  in  woods,  rare.  Bemidji,  July  n,  1900,  no.  645. 
Thief  River  Falls,  July  17,  1900,  no.  752. 

Spores  occasionally  more  than  four-celled  and  frequently 
narrower  than  usual. 

45.  Peltigera  refuscens  (NECK.)  HOFFM. 

On  earth,  rare  or  infrequent.  Battle  lake,  June  21,  1900,  no. 
118.  Henning,  June  30,  1900,  no.  372.  Thief  River  Falls, 
July  21,  1900,  no.  862. 

46.  Peltigera  canina  (L.)  HOFFM. 

On  earth,  common.  Battle  lake,  June  18,  1900,  no.  14  and 
June  20,  1900,  no.  81.  Henning,  June  25,  1900,  no.  186. 
Leaf  hills,  June  26,  1900,  no.  237.  Thief  River  Falls,  July 
17,  1900,  no.  751.  Red  lake,  July  27,  1900,  no.  933. 

47.  Peltigera  canina  (L.)  HOFFM.  var.  spuria  ACH. 

On  earth,  frequent.  Battle  lake,  June  20,  1900,  no.  69. 
Leaf  hills,  June  26,  1900,  no.  223.  Bemidji,  July  9,  1900, 
nos.  533  and  545.  Thief  River  Falls,  July  19,  1900,  no.  809. 
Red  lake,  July  27,  1900,  no.  909. 


Fink:   LICHENS  OF  NORTHWESTERN  MINNESOTA.  689 

48.  Peltigera  canina  (L.)  HOFFM.  var.  sorediata  SCHAER. 

On  earth,  frequent.  Battle  lake,  June  23,  1900,  no.  152. 
Leaf  hills,  June  26,  1900,  no.  216.  Bemidji,  July  4,  1900, 
no.  439.  Thief  River  Falls,  July  17,  1900,  no.  765.  Red 
lake,  July  27,  1900,  nos.  921  and  931. 

49.  Peltigera  canina  (L.)  HOFFM.  var.  leucorrhiza  FLK. 

On  earth  in  tamarack  swamps,  infrequent.  Henning,  June 
25,  1900,  no.  194  and  June  29,  1900,  no.  342.  Bemidji,  July 
9,  1900,  nos.  534  and  546.  Red  lake,  July  28,  1900,  no.  941. 

Not  previously  reported  from  Minnesota. 

50.  Heppia  despreauxii  (MONT.)  TUCK. 

On  calcareous  earth,  infrequent.  Battle  lake,  June  20, 
1900,  no.  57.  Leaf  hills,  June  26,  1900,  no.  232,  June  27, 
1900,  no.  264  and  July  2,  1900,  no.  394.  Thief  River  Falls, 
July  17,  1900,  no.  773. 

51.  Pannaria  languinosa  (AcH.)  KBR. 

On  mossy  tree  bases,  rare.  Henning,  June  28,  1900,  no. 
300.  Bemidji,  July  5,  1900,  no.  458,  July  9,  1900,  no.  549 
and  July  n,  1900,  no.  625.  Thief  River  Falls,  July  17,  1900, 
no.  763.  Red  lake,  July  28,  1900,  no.  946. 

52.  Pannaria  petersii  TUCK. 

On  calcareous  pebbles,  rare.  Leaf  hills,  July  2,  1900,  no. 
399.  With  thallus  nearly  obsolete,  the  same  condition  occur- 
ring occasionally  in  northern  Iowa. 

Not  previously  reported  from  Minnesota. 

53.  Collema  pycnocarpum  NYL. 

On  trees,  rare.  Battle  lake,  June  19,  1900,  no.  33  and  June 
21,  1900,  no.  no.  Bemidji,  July  5,  1900,  no.  467,  July  n, 
1900,  no.  607,  July  12,  1900,  no.  686  and  July  14,  1900,  no. 
724.  Red  lake,  August  3,  1900,  no.  1069. 

54.  Collema  flaccidum  ACH. 

On  trees,  infrequent.  Battle  lake,  June  23,  1900,  no.  155. 
Henning,  June  29,  1900,  no.  351.  Bemidji,  July  12,  1900,  no. 
668. 

55.  Collema  nigrescens  (Huos.)  ACH. 

On  trees,  frequent.  Henning,  June  29,  1900,  nos.  349  and 
359.  Bemidji,  July  12,  1900,  no.  693.  Red  lake,  August  3, 
1900,  no.  1074. 


690  MINNESOTA    BOTANICAL    STUDIES. 

56.  Collema  ryssoleum  TUCK. 

On  Popttht3t  rare.      Bemidji,  July   12,   1900,   no.  686. 
Not  previously  reported  from  Minnesota. 

57.  Collema  pulposum  (BERNH.)  NYL. 

On  earth,  rare.  Battle  lake,  June  20,  1900,  no.  77.  Leaf 
hills,  June  26,  1900,  no.  207.  Henning,  June  29,  1900,  no. 
352.  Leaf  hills,  July  2,  1900,  no.  400.  Bemidji,  July  5, 
1900,  no.  477.  Thief  River  Falls,  July  17,  1900,  nos.  769  and 
770. 

58.  Collema  crispum  BORR. 

On  earth,  frequent.     Bemidji,  July  n,  1900,  no.  605. 

59.  Collema  limosum  ACH. 

On    earth,   infrequent.     Bemidji,    July    10,    1900,    no.    568, 
July  n,  1900,  no.  637  and  July  14,  1900,  no.  736. 
Not  previously  reported  from  Minnesota. 

60.  Leptogium  lacerum  (Sw.)  FR. 

On  earth,  rare.  Leaf  hills,  June  27,  1900,  no.  287.  Thief 
River  Falls,  July  18,  1900,  no.  795. 

61.  Leptogium  myochroum  (EHRH.,  SCHAER.)  TUCK. 

On  trees  and  rocks,  rare.  Battle  lake,  June  21,  1900,  no. 
123  and  June  23,  1900,  no.  156.  Henning,  June  30,  1900,  nos. 
381  and  382.  Bemidji,  July  5,  1900,  no.  485  and  July  13, 
1900,  no.  710. 

62.  Placodium  elegans  (LINK.)  DC. 

On  granite  and  limestone,  rare.  Battle  lake,  June  20,  1900, 
nos.  60  and  66.  Bemidji,  July  n,  1900,  no.  638.  Thief  River 
Falls,  July  17,  1900,  no.  776. 

63.  Placodium  murorum  (HOFFM.)  DC. 

On  lime  bowlders,  rare.  Thief  River  Falls,  July  9,  1900, 
no.  825.  With  deficient  thallus. 

64.  Placodium  cinnabarrinum  (Acn.)  ANZ. 

On  granite  rocks,  infrequent.  Battle  lake,  June  20,  1900, 
nos.  45  and  50.  Leaf  hills,  June  27,  1900,  nos.  276  and  282. 

65.  Placodium  aurantiacum  (LIGHTF.)  NAEG.  and  HEPP. 

On  trees  and  granites,  infrequent.  Battle  lake,  June  19, 
1900,  no.  37.  Leaf  hills,  June  27,  1900,  no.  253.  Bemidji, 
July  n,  1900,  no.  643.  Thief  River  Falls,  July  19,  1900,  no. 


Fink:    LICHENS  OF  NORTH\VKSTEK\   MINNESOTA.  091 

819    and   July    21,    1900,    no.    855.      Some    have   very   heavy 
thallus. 

66.  Placodium  cerinum   (HEDW.)  NAEG.  and  HEPP. 

On  trees  and  granite  rocks,  frequent  or  infrequent.  Battle 
lake,  June  18,  1900,  no.  13,  June  19,  1900,  no.  30,  June  20, 
1900,  no.  73  and  June  23,  1900,  no.  164.  Henning,  June  25, 
1900,  nos.  177  and  199.  Leaf  hills,  June  26,  1900,  no.  244 
and  June  27,  1900,  nos.  267  and  271.  Henning,  June  28, 
1900,  no.  321.  Bemidji,  July  4,  1900,  no.  454,  July  5,  1900, 
nos.  459  and  465  and  July  n,  1900,  no.  644.  Thief  River 
Falls,  July  17,  1900,  nos.  756  and  775,  July  19,  1900,  nos. 
817,  818  and  823,  July  20,  1900,  no.  835  and  July  21,  1900, 
no.  861.  Red  lake,  July  26,  1900,  no.  883,  July  28,  1900, 
no.  953,  July  30,  1900,  no.  979,  Aug.  i,  1900,  no.  1025  and 
Aug.  2,  1900,  no.  1061. 

67.  Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP.  var.  sidiritis 
TUCK. 

On  granite  rocks,  infrequent.  Battle  lake,  June  20,  1900, 
no.  68,  and  June  23,  1900,  nos.  149  and  157.  Leaf  hills,  June 
27,  1900,  nos.  247  and  263.  Red  lake,  July  31,  1900,  no. 
ion. 

68.  Placodium  cerinum  (HEDW.)  NAEG.  and  HEPP.  var.  pyracea 
NYL. 

On  pine  logs,  frequent.     Red  lake,  July  30,  1900,  no.  983. 

69.  Placodium  ferrugineum  (Huos.)  HEPP. 

On  old  pine,  frequent.  Bemidji,  July  15,  1900,  no.  705. 
Red  lake,  July  30,  1900,  no.  976. 

70.  Placodium  vitellinum  (EHRH.)  NAEG.  and  HEPP. 

On  old  wood  and  granite  rocks,  rare.  Battle  lake,  June  20, 
1900,  no.  104.  Leaf  hills,  June  26,  1900,  no.  240  and  June 
27,  1900,  no.  278.  Bemidji,  July  10,  1900,  no.  559.  Red 
lake,  July  28,  1900,  no.  960  and  July  30,  1900,  no.  982. 

71.  Placodium    vitellinum  (EHRH.)  NAEG.    and    HEPP.    var. 
aurellum  ACH. 

On  trees  and  granite  rocks,  infrequent.  Battle  lake,  June 
20,  1900,  no.  56.  Leaf  hills,  June  26,  1900,  no.  25 la  and  June 
27,  1900,  no.  257.  Bemidji,  July  u,  1900,  no.  639. 


692  MINNESOTA    BOTANICAL    STUDIES. 

72.  Lecanora  rubina  (VILL.)  ACH. 

On  granitic  rocks,  infrequent  or  rare.  Battle  lake,  June  20, 
1900,  nos.  46,  49  and  51.  Leaf  hills,  June  27,  1900,  no.  284. 
Bemidji,  July  u,  1900,  no.  594. 

73.  Lecanora  muralis  (SCHREB.)  SCHAER. 

On  rocks,  rare.     Red  lake,  August  i,  1900,  no.  1030. 

74.  Lecanora  muralis  (SCHREB.)  SCHAER.  var.  saxicola  SCHAER. 
On  granitic  rocks,  locally  frequent.      Battle   lake,  June  20, 

1900,  no.  107.     Leaf  hills,  June  26,  1900,  no.  201. 

75.  Lecanora  muralis  (SCHREB.)  SCHAER.  var.  versicolor  FR. 
On    lime    rocks,  locally   frequent.       Battle    lake,  June    20, 

1900,  nos.  83,  97  and  102. 

76.  Lecanora  pallida  (SCHREB.)  SCHAER. 

On  cedar  in  swamp,  rare.     Bemidji,  July  13,  1900,  no.  717. 

77.  Lecanora  subfusca  (L.)  ACH. 

On  trees  and  rocks,  common  in  last  two  localities  only, 
Battle  lake,  June  20,  1900,  nos.  54  and  75.  Henning,  June 
25,  1900,  no.  198.  Leaf  hills,  June  27,  1900,  no.  279. 
Bemidji,  July  4,  1900,  no.  457,  July  5,  1900,  nos.  471  and  499 
and  July  u,  1900,  no.  606.  Thief  River  Falls,  July  20,  1900, 
no.  839.  Red  lake,  August  3,  1900,  no.  1075. 

No.  54  is  a  peculiar  form  on  rocks  with  scant  thallus.  No. 
606  has  border  of  exciple  much  raised. 

78.  Lecanora  subfusca  (L.)  ACH.  var.  argentata  ACH. 
On  pines,  rare.     Red  lake,  August  3,  1900,  no.  1066. 

79.  Lecanora  subfusca  (L.)  ACH.  var.  coilocarpa  ACH. 
On  pine  logs,  rare.     Red  lake,  July  30,  1900,  no.  993. 

80.  Lecanora  variolascens  NYL. 

On  trees,  common  in  Minnesota  and  Iowa,  but  usually  sterile 
and  hence  not  determinable.  Battle  lake,  June  20,  1900,  no. 
76.  Henning,  June  29,  1900,  no.  358.  Thief  River  Falls, 
July  18,  1900,  no.  800.  Red  lake,  July  28,  1900,  no.  975. 

Not  previously  reported  from  Minnesota  and  new  to  North 
America. 

81.  Lecanora  hageni  ACH. 

On  rocks,  rare.  Battle  lake,  June  20,  1900,  no.  109. 
Bemidji,  July  IT,  1900,  no.  640. 


Fink:    LICHENS  OF  NORTHWESTERN  MINNESOTA.  693 

82.  Lecanora  varia  (EHRH.)  NYL. 

On  trees,  dead  wood  and  rocks,  infrequent.  Battle  lake, 
June  23,  1900,  nos.  136  and  139.  Henning,  June  25,  1900, 
no.  184.  Leaf  hills,  June  27,  1900,  no.  252.  Henning,  June 
28,  1900,  nos.  309  and  333.  Bemidji,  July  5,  1900,  nos.  492, 
498  and  500  and  July  12,  1900,  nos.  695  and  697.  Thief  River 
Falls,  July  17,  1900,  no.  758  and  July  18,  1900,  nos.  784,  785 
and  804.  Red  lake,  July  27,  1900,  no.  919  and  August  I, 
1900,  no.  1022. 

83.  Lecanora  varia  (EHRH.)  NYL.  var.  polytropa  NYL. 

On  granitic  rocks,  rare.       Battle  lake,  June  20,   1900,   no. 

84.  Thief  River  Falls,  July  21,   1900,   no.  864.     Red  lake, 
July  27,  1900,  no.  905. 

Not  previously  reported  from  Minnesota. 

84.  Lecanora  varia  (EHRH.)  NYL.  var.  symmicta  ACH. 

On  dead  wood,  rare  but  widely  distributed.  Battle  lake, 
June  23,  1900,  nos.  138  and  142.  Henning,  June  28,  1900, 
no.  336.  Bemidji,  July  6,  1900,  no.  545.  Thief  River  Falls, 
July  18,  1900,  no.  806  and  July  21,  1900,  no.  867.  Red  lake, 
July  28,  1900,  nos.  954  and  966  and  August  2,  1900,  no.  1060. 

85.  Lecanora  varia  (EHRH.)  NYL.  var.  saspincola  FR. 

On  old  wood,  rare.  Battle  lake,  June  23,  1900,  no.  150. 
Red  lake,  July  28,  1900,  no.  957. 

86.  Lecanora  pallescens  (L.)  SCHAER. 

On  trees  in  swamp,  rare.     Bemidji,  July  13,  1900,  no.  708. 

87.  Lecanora  verrucosa  (Acn.)  LAUR.  var.  mutabilis  TH.  FR. 
On  trees,  rare.      Henning,  June  29,  1900,  no.  350.     Bemidji, 

July  n,  1900,  no.  609,  July  13,  1900,  no.  713  and  July  14,  1900, 
no-  735-     R-ed  lake,  August  3,  1900,  no.  1067. 

Not  previously  reported  from  Minnesota,  but  confused  with 
Pertusaria  leioplaca  (Ach.)  Schaer. 

88.  Lecanora  cinerea  (L.)  SOMMERF. 

On  granitic  bowlders,  frequent.  Battle  lake,  June  20,  1900, 
nos.  91,  99  and  101.  Leaf  hills,  June  26,  1900,  no.  233  and 
June  27,  1900,  nos.  249,  260  and  295.  Bemidji,  July  10,  1900, 
no.  573,  July  n,  1900,  nos.  592  and  599  and  July  14,  1900, 
no.  737. 


694  MINNESOTA    BOTANICAL    STUDIES. 

89.  Lecanora  calcarea  (L.)  SOMMERF.  var.  contorta  FR. 

On  bowlders,  especially  calcareous,  rare.  Battle  lake,  June 
20,  1900,  nos.  48,  92  and  100.  Leaf  hills,  June  27,  1900,  no. 
296. 

90.  Lecanora  gibbosa  (Acn.)  NYL.  var  microspora  A.  ZAHLB. 
On  exposed  granitic  bowlders,  frequent.     Leaf  hills,  July  2, 

1900,  no.  388. 

Not  previously  reported  from  Minnesota  and  new  to  North 
America.  Synonymy  uncertain. 

91.  Lecanora  cervina  (PERS.)  NYL. 

On  granitic  bowlders,  infrequent.  Leaf  hills,  June  26,  1900, 
nos.  205  and  238,  June  27,  1900,  no.  251  and  July  2,  1900,  no. 
397.  Bemidji,  July  16,  1900,  no.  744. 

92.  Lecanora  fuscata  (SCHRAD.)  TH.  FR. 

On  granitic  bowlders,  infrequent.  Battle  lake,  June  20, 
1900,  no.  80.  Leaf  hills,  June  26,  1900.  no.  218  and  June 
27,  1900,  no.  280.  Bemidji,  July  10,  1900,  no.  560.  Red 
lake,  July  27,  1900,  no.  924. 

93.  Lecanora  privigna  (Acn.)  NYL. 

On  bowlders,  especially  calcareous,  rare.  Leaf  hills,  June 
27,  1900,  nos.  259,  288  and  292  and  July  2,  1900,  no.  393. 
Bemidji,  July  10,  1900,  no.  584. 

The  last  is  a  peculiar  clustered  form  on  granitic  bowlders. 

94.  Lecanora  privigna  (Acn.)  NYL.  var.  pruinosa  AUCT. 

On  calcareous  bowlders,  rare.  Leaf  hills,  July  2,  1900,  no. 
404. 

95.  Lecanora  xanthophana  NYL. 

On  granitic  bowlders,  locally  frequent  on  high  hills.  Battle 
lake,  June  20,  1900,  no.  47  and  June  23,  1900,  no.  167. 

96.  Rinodina  oreina  (Acn.)  MASS. 

On  granitic  rocks,  common  in  first  locality.  Battle  lake, 
June  20,  1900,  nos.  90  and  106.  Leaf  hills,  June  26,  1900, 
no.  202.  Thief  River  Falls,  July  21,  1900,  no.  865.  Red 
lake,  August  i,  1900,  no.  1021. 

97.  Rinodina  sophodes  (Acn.)  NYL. 

On  trees,  old  wood  and  rocks,  common.  Battle  lake,  June 
18,  1900,  no.  19,  June  21,  1900,  no.  119  and  June  23,  1900, 
no.  162.  Leaf  hills,  June  26,  1900,  nos.  222  and  231.  Hen- 


Fink :   LICHENS  OF  NORTHWESTERN  MINNESOTA.  695 

ning,  June  28,  1900,  no.  339.  Bemidji,  July  5,  1900,  nos.  464 
and  478,  July  7,  1900,  no.  529  and  July  n,  1900,  no.  651. 
Thief  River  Falls,  July  18,  1900,  no.  808,  July  19,  1900,  no. 
811  and  July  21,  1900,  no.  847.  Red  lake,  July  26,  no.  888 
and  July  28,  1900,  nos.  935  and  969. 

Spores  commonly  smaller  than  Tuckerman's  measurements. 

98.  Rinodina  sophodes  (AcH.)  NYL.  var.  atrocinerea  NYL. 

On  tamarack  in  swamps,  rare.  Bemidji,  July  6,  1900,  nos. 
538  and  547. 

Not  previously  reported  from  Minnesota. 

99.  Rinodina  sophodes  (Acn.)  NYL.  var.  exigua  FR. 

On  granitic  rocks,  rare.  Battle  lake,  June  23,  1900,  no. 
153.  Leaf  hills,  June  26,  1900,  no.  217. 

100.  Rinodina  bischoffi  (HEPP.)  KBR. 

On  calcareous  rocks,  rare.  Leaf  hills,  June  27,  1900,  no. 
291. 

101.  Rinodina  nigra  sp.  nov. 

On  granitic  rocks,  infrequent.  Battle  lake,  June  23,  1900, 
no.  146. 

Thallus  thin  (12-22  mm.),  indeterminate  or  subdeterminate, 
tartareous-areolate,  ecorticate ;  the  areoles  densely  crowded, 
angular,  .5-1.25  mm.  wide,  dark  slate  color;  upon  a  black 
hypothallus,  which  also  borders  the  thallus  more  or  less ;  algal 
cells  globose,  green  (Cystococcus?),  8-14  mic.  in  diameter. 
Apothecia  minute,  . 25-^0  mic.  in  diameter,  immersed  in  the 
thallus,  one,  two  or  three  in  each  areole  ;  disk  black,  usually 
somewhat  depressed,  circular,  with  an  entire  thalline  exciple, 
concolorous  with  the  thallus ;  hymenium  75—90  mic.  deep. 
Spores  brown,  2-celled,  oblong  and  frequently  somewhat  con- 
stricted at  the  septum,  9-15  mic.  long  and  5-8  mic.  wide,  8  in 
asci.  Paraphyses  distinct,  slender,  unbranched,  colorless 
throughout  or  brownish  at  tips.  Asci  50-60  by  13-17  mic. 

102.  Pertusaria  multipuncta  (TURN.)  NYL. 

On  trees,  frequent.  Bemidji,  July  13,  1900,  no.  711  and 
July  14,  1900,  nos.  723,  725  and  726. 

103.  Pertusaria  communis  DC. 

On  trees  in  swamp,  frequent.  Bemidji,  July  13,  1900,  nos. 
702  and  704. 


696  MINNESOTA    BOTANICAL    STUDIES. 

104.  Pertusaria  pustulata  (Acn.)  NYL. 

On, trees,  infrequent.  Battle  lake,  June  22,  1900,  no.  129. 
Henning,  June  29,  1900,  no.  361  and  June  30,  1900,  no.  383. 
Bemidji,  July  13,  1900,  nos.  707  and  719. 

105.  Pertusaria  finkii  A.  ZAHLB.  in  litt. 

On  trees,  rare  or  infrequent.  Battle  lake,  June  20,  1900, 
no.  74,  June  21,  1900,  no.  113  and  June  26,  1900,  no.  128. 
Henning,  June  30,  1900,  no.  366.  Bemidji,  July  5,  1900,  no. 
480.  Thief  River  Falls,  July  18,  1900,  no.  801.  Red  lake, 
July  31,  1900,  nos.  1014  and  1019. 

Thallus  white  or  whitish,  thin  (0.2-0.28  mm.),  determinate  or 
subdeterminate,  margin  continuous,  center  unequal-verrucose 
and  rimose,  ecorticate,  medullary  hyphge  slender,  algal  cells 
globose,  green,  10—15  m^c.  ^n  diameter,  disposed  in  more  or  less 
discrete  clusters.  Apothecia  plentiful,  clustered,  immersed  or 
finally  becoming  open-lecanoroid,  about  0.75  mm.  in  diameter, 
irregular  or  subrotund,  fuscescent  or  blackish,  flat,  rough ; 
thalline  margin  persistent,  thin,  whitish,  crenulate  to  sublacerate. 
Hymenium  320-370  mic.  deep.  Epithecium  rufescent  or  fus- 
cescent. Paraphyses  slender,  somewhat  conglutinate.  Asci 
elevate-saccate,  containing  2  (rarely  i)  spores,  straight  or  some- 
what curved,  apex  rounded  and  thickened,  170-190  mic.  long 
and  42-46  mic.  wide.  Spores  oval,  ellipsoid  or  oblong,  occa- 
sionally somewhat  constricted  along  the  sides,  simple,  colorless, 
88-140  mic.  long  and  28-53  rrnc-  wide,  spore  walls  thick  (9- 
12  mic.). 

1 06.  Urceolaria  scruposa  (L.)  NYL. 

On  calcareous  earth,  rare.  Battle  lake,  June  20,  1900,  nos. 
52  and  96.  Leaf  hills,  June  27,  1900,  no.  255. 

107.  Stereocaulon  paschale  (L.)  FR. 

On  granitic  bowlders,  rare.  Bemidji,  July  10,  1900,  no. 
569  and  July  16,  1900,  no.  750.  Red  lake,  August  4,  1900, 
nos.  1079  and  1082. 

1 08.  Cladonia  mitrula  TUCK. 

On  earth,  rare.  Battle  lake,  June  20,  1900,  no.  53.  Hen- 
ning, June  28,  1900,  no.  308.  Leaf  hills,  July  2,  1900,  no.  402. 

109.  Cladonia  botrytes  (HAG.)  WILLD. 

\.  ,  On  coniferous  stumps  and  logs,  infrequent.  Bemidji,  July 
4,  1900,  no.  430  and  July  6,  1900,  no.  526.  Thief  River 
Falls,  July  17,  1900,  no.  771.  Red  lake,  July  28,  1900,  no. 

974- 


Fink:   LICHENS  OF  NORTHWESTERN  MINNESOTA.  697 

Not  previously  reported  from  Minnesota. 

no.  Cladonia  cariosa  (Acn.)  SPRENG. 

On  earth,  infrequent.  Leaf  hills,  June  26,  1900,  nos.  226, 
227  and  229.  Bemidji,  July  7,  1900,  no.  503,  July  10,  1900, 
nos.  565  and' 566  and  July  16,  1900,  no.  741.  Red  lake, 
August  3,  1900,  nos.  1075  and  1080. 

in.  Cladonia  pyxidata  (L.)  FR. 

On  earth  and  old  logs,  frequent.  Henning,  June  25,  1900, 
no.  180  and  June  27,  1900,  no.  255.  Thief  River  Falls,  July 
17,  1900,  no.  766.  Red  lake,  July  27,  1900,  no.  907. 

112.  Cladonia  pyxidata  (L.)  FR.  var.  neglecta  (FLK.)  MASS. 
On  old  logs,  frequent.     Battle  lake,  June  18,  1900,  no.  12. 

Thief  River  Falls,  July  17,  1900,  no.  757.     Red  lake,  August 
2,  1900,  no.  1055. 

Not  previously  reported  from  Minnesota. 

113.  Cladonia  pyxidata  (L.)  FR.  var.  chlorophaea  FLK. 

On  old  stumps,  infrequent.     Bemidji,  July  6,  1900,  no.  519. 
Not  previously  reported  from  Minnesota. 

114.  Cladonia  fimbriata  (L.)  FR.  var.  tubaeformis  FR. 

On  old  wood,  common.  Battle  lake,  June  20,  1900,  no. 
78.  Henning,  June  25,  1900,  no.  185  and  June  28,  1900,  no. 
306.  Bemidji,  July  4,  1900,  no.  438  and  July  7,  1900,  nos. 

551  and  552. 

115.  Cladonia  fimbriata  (L.)  FR.  var.  simplex  (WEIS.)  FLOT. 
On    rotten   wood,    rare.      Henning,  July   2,    1900,    no    407. 

Bemidji,  July  6,   1900,   no.  533.     Apparently  a  synomym  for 
above  according  to  Wainio,  but  specimens  very  different. 
Not  previously  reported  from  Minnesota. 

116.  Cladonia  fimbriata  (L.)  FR.  var.  subulata  (L.)  WAINIO. 
On  earth  in  pine  woods,  infrequent.     Bemidji,  July  10,  1900, 

no.  574.     Red  lake,  July  26,  1900,  no.  902. 
Not  previously  reported  from  Minnesota. 

117.  Cladonia  fimbriata  (L.)  FR.  var.  fibula  ACH. 

On  earth  usually  under  conifers,  infrequent.  Leaf  hills, 
June  26,  1900,  no.  230.  Bemidji,  July  5,  1900,  nos.  462,  504 
and  509,  July  9,  1900,  no.  535  and  July  10,  1900,  nos.  564,  571 
and  585.  Red  lake,  August  3,  1900,  no.  1076. 

Not  previously  reported  from  Minnesota. 


698  MINNESOTA    BOTANICAL    STUDIES. 

118.  Cladonia  fimbriata  (L.)  FR.  var.  apolepta  (Acn.)  WAINIO. 
On  old  wood,  especially  tamarack,  frequent.      Henning,  June 

28,  1900,  no.  303.    Bemidji,  July  6,  1900,  no.  521.    Thief  River 
Falls,  July  19,  1900,  no.  815. 

Not  previously  reported  from  Minnesota  and  new  to  North 
America. 

119.  Cladonia  fimbriata  (L.)  FLK.  var.  ceratodes  (FLK.)  WAINIO. 
On  earth  under  pines,  common.     Bemidji,  July  6,  1900,  no. 

486.      Red  lake,  July  28,  1900,  no.  950. 

Not  previously  reported  from  Minnesota  and  new  to  North 
America. 

1 20.  Cladonia  degenerans  FLK. 

On  earth  under  pines,  rare.      Bemidji,  July  5,  1900,  no.  510. 

121.  Cladonia  gracilis  (L.)  NYL. 

On  earth  and  old  wood,  frequent.  Battle  lake,  June  20, 
1900,  no.  79.  Henning,  June  28,  1900,  nos.  311  and  319. 
Bemidji,  July  4,  1900,  no.  431.  Red  lake,  July  30,  1900,  no. 
987. 

122.  Cladonia  gracilis  (L.)  NYL.  var.  verticellata  FR. 

On  earth  and  old  wood,  frequent.  Battle  lake,  June  18, 
1900,  no.  ii.  Leaf  hills,  June  26,  1900,  nos.  219,  220  and 
221.  Henning,  June  28,  1900,  no.  320.  Bemidji,  July  5, 
1900,  no.  572  and  July  14,  1900,  no.  732.  Red  lake,  July 

27,  1900,  nos.  922  and  934&. 

123.  Cladonia  gracilis  (L.)  NYL.  var.  anthocephala  FLK. 

On   old   logs,  usually  coniferous,  frequent.     Henning,  June 

28,  1900,  no.   314.     Bemidji,  July  6,  1900,  nos.   524  and  528 
and  July  10,  1900,  no.  582.     Thief  River  Falls,  July  19,  1900, 
no.  797.     Red  lake,  July  28,  1900,  no.  968. 

Not  previously  reported  from  Minnesota  and  new  to  North 
America. 

124.  Cladonia  gracilis  (L.)  NYL.  var.  hybrida  SCHAER. 

On  earth  and  old  logs,  common.  Henning,  June  28,  1900, 
nos.  302  and  304.  Bemidji,  July  4,  1900,  no.  437,  July  5, 
1900,  nos.  507  and  508  and  July  6,  1900,  nos.  515  and  550. 
Red  lake,  July  27,  1900,  no.  934. 

125.  Cladonia  cenotea  (Acn.)  SCHAER. 

On  earth,  rare.     Bemidji,  July  9,  1900,  nos.  534  and  536. 


Fink:     LICHENS    OF    NORTHWESTERN    MINNESOTA.  699 

126.  Cladonia  squamosa  HOFFM. 

On  old  stumps  in  tamarack  swamp,  locally  common.  Be- 
midji,  July  6,  1900,  no.  516. 

127.  Cladonia  furcata  (HUBS.)  FR. 

On  earth,  rare.  Bemidji,  July  12,  1900,  no.  684  and  July 
13,  1900,  no.  720. 

128.  Cladonia  furcata (HuDs.)FR.var.  scabriuscula(DEL.)CEON. 
On  earth  or  old  logs,  rare.      Henning,  July  2,  1900,  no.  416. 

Bemidji,  July  13,  1900,  no.  720. 

Not  previously  reported  from  Minnesota  and  new  to  North 
America. 

129.  Cladonia  furcata  (HUDS.)  FR.  var.  paradoxa  WAINIO. 

On  earth  or  old  wood  under  conifers,  frequent.  •  Henning, 
June  30,  1900,  no.  371.  Bemidji,  July  4,  1900,  no.  432,  July 

6,  1900,  no.   522,  July  10,  1900,   nos.   562,  570,  567,   and  586 
and  July  12,  1900,  no.  677.     Red  lake,  July  27,  1900,  no.  906. 

Not  previously  reported  from  Minnesota  and  new  to  North 
America. 

130.  Cladonia  rangiferina  (L.)  HOFFM. 

On  earth  and  logs,  common  under  pines  in  second  locality. 
Henning,  June  25,  1900,  no.  181,  June  28,  1900,  no.  327  and 
June  29,  1900,  no.  340.  Bemidji,  July  5,  1900,  no.  505,  July 

7,  1900,  no.  509,  July  10,  1900,  no.  587  and  July  14,  1900,  nos. 
729  and  738.     Thief  River  Falls,  July  17,  1900,  no.  761.     Red 
lake,  July  26,  1900,  no.  903  and  July  27,  1900,  no.  923. 

131.  Cladonia  rangiferina  (L.)  HOFFM.  var.  sylvatica  (L.)  RA- 
BENH. 

On  earth  under  conifers,  common.  Bemidji,  July  5,  1900, 
nos.  503  and  511,  July  6,  1900,  no.  517  and  July  14,  1900,  no. 
739.  Red  lake,  July  27,  1900,  nos.  908  and  915. 

132.  Cladonia  rangiferina  (L.)  HOFFM.  var.  alpestris  L. 

On  earth,  infrequent.  Bemidji,  July  6,  1900,  no.  517  and 
July  14,  1900,  no.  739.  Red  lake,  July  27,  1900,  no.  915. 

133.  Cladonia  uncialis  (L.)  FR. 

On  earth  in  pine  woods,  rare.  Bemidji,  July  9,  1900,  no. 
538. 

134.  Cladonia  deformis  (L.)  HOFFM. 

On  old  logs,  rare.     Bemidji,  July  14,  1900,  no.  734. 


700  MINNESOTA    BOTANICAL    STUDIES. 

135.  Cladonia  digitata  (L.)  HOFFM. 

On  tamarack  log  in  swamp,  rare.      Bemidji,  July  6,  1900,  no. 


136.  Cladonia  bacillaris  NYL. 

On  old  logs  under  pines,  infrequent.  Bemidji,  July  4,  1900, 
no.  419  and  July  9,  1900,  nos.  540  and  541. 

Not  previously  reported  from  Minnesota,  but  included  under 
the  next. 

137.  Cladonia  macilenta  (EHRH.)  HOFFM. 

On  old  logs,  infrequent.  Bemidji,  July  9,  1900,  no.  540. 
Red  lake,  July  28,  1900,  no.  937. 

138.  Cladonia  cristatella  TUCK. 

On  old  wood  and  earth,  common.  Battle  lake,  June  20, 
1900,  nos.  41  and  42.  Leaf  hills,  June  26,  1900,  no.  225. 
Henning,  June  28,  1900,  nos.  422  and  433,  July  7,  1900,  no. 
435  »  JulJ  8»  1900,  no-  474>  July  9>  I9°°>  nos.  537  and  549  and 
July  10,  1900,  no.  577.  Thief  River  Falls,  July  18,  1900,  no. 
778.  Red  lake,  July  27,  1900,  no.  904  and  July  28,  1900,  no. 
956. 

139.  Biatora  decipiens  (EHRH.)  FR. 

On  calcareous  earth,  frequent.  Battle  lake,  June  20,  1900, 
no.  59.  Leaf  hills,  June  27,  1900,  no.  275  and  July  2,  1900, 
no.  391. 

140.  Biatora  decipiens  (EHRH.)  FR.  var.  dealbata  TUCK. 

On  earth,  infrequent.  Leaf  hills,  June  27,  1900,  no.  261  and 
July  2,  1900,  no.  392. 

141.  Biatora  viridescens  (SCHRAD.)  FR. 

On  tamarack  logs  in  swamp,  frequent.  Bemidji,  July  7, 
1900,  no.  528. 

142.  Biatora  vernalis  (L.)  FR. 

On  mossy  trees  and  on  tamarack,  infrequent.  Bemidji,  July 
7,  1900,  nos.  506,  508,  527  and  537  and  July  13,  1900,  no.  718. 

143.  Biatora  turgidula  (FR.)  NYL, 

On  elms,  rare.     Bemidji,  July  9,  1900,  no.  553. 

144.  Biatora  leucophaea  (FLK.)  TUCK. 

On  granite  rocks,  infrequent.  Leaf  hills,  June  27,  1900,  no. 
277.  Bemidji,  July  n,  1900,  no.  602.  Thief  River  Falls, 


Fink:     LICHENS    OF    NORTHWESTERN    MINNESOTA.  701 

July  18,  1900,  no.  792.      Red  lake,  August  i,  1900,  nos.  1031 
and  1042. 

145.  Biatora  uliginosa  (SCHRAD.)  FR. 

On  pine  logs  or  earth  under  pines,  infrequent.  Bemidji, 
July  4,  1900,  no.  440,  July  5,  1900,  no.  487  and  July  9,  1900, 
nos.  539  and  542.  Red  lake,  July  28,  1900,  no.  971,  July  30, 
1900,  nos.  977  and  Aug.  i,  1900,  no.  1046. 

146.  Biatora  myriocarpoides  (NYL.)  TUCK. 

On  old  pine  and  granite,  infrequent.  Bemidji,  July  4,  1900, 
no.  424,  July  10,  1900,  no.  561  and  July  16,  1900,  nos.  743 
and  749.  Thief  River  Falls,  July  20,  1900,  no.  840,  July  21, 
1900,  no.  851  and  July  23,  1900,  nos.  870  and  873.  Red  lake, 
July  28,  1900,  nos.  959  and  967,  July  30,  1900,  no.  1002,  Aug. 
2,  1900,  no  1058  and  Aug.  4,  1900,  no.  1077. 

Part  of  the  material  here  referred  to  has  a  colorless  hypothe- 
cium  and  will  doubtless  be  referred  elsewhere  after  further 
study. 

147.  Biatora  varians  (Acn.)  TUCK. 

On  trees,  infrequent.  Battle  lake,  June  18,  1900,  no.  i  and 
June  23,  1900,  no.  163.  Leaf  hills,  June  26,  1900,  no.  214. 
Henning,  June  29,  1900,  no.  356.  Thief  River  Falls,  July  21, 
1900,  no.  860. 

148.  Biatora  mixta  FR. 

On  poplars,  frequent.  Henning,  June  29,  1900,  no.  434. 
Bemidji,  July  n,  1900,  no.  629.  Thief  River  Falls,  July  21, 
1900,  no.  854.  Red  lake,  Aug.  2,  1900,  no.  1062. 

Not  previously  reported  from  Minnesota. 

149.  Biatora  mixta  FR.  var.  atlantica  TUCK. 

On  poplars,  rare.     Battle  lake,  June  20,  1900,  no.  70. 
Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

150.  Biatora  atropurpurea  (MASS.)  HEPP. 

On  poplars,  infrequent.  Bemidji,  July  4,  1900,  no.  455  and 
July  9,  1900,  no.  551,  Thief  River  Falls,  July  21,  1900,  nos. 
777  and  846. 

151.  Biatora  prasina  FR.  var.  byssacea  TH.  FR. 

On  old  logs  in  swamps,  rare.  Bemidji,  July  7,  1900,  no. 
520. 

Not  previously  reported  from  Minnesota. 


702  MINNESOTA    BOTANICAL    STUDIES. 

152.  Biatora  glauco-nigrans  TUCK. 

On  trees  in  low  woods,  infrequent.      Henning,  June  29,  1900, 
no.  355- 

153.  Biatora  sphaeroides  (DICKS.)  TUCK. 

On  mossy  bases,  rare.     Bemidji,  July  12,  1900,  no.  663. 
Not  previously  reported  from  Minnesota. 

154.  Biatora  hypnophila  (TURN.)  TUCK. 

On  mossy  earth,  rare.  Henning,  June  29,  1900,  no.  346 
and  June  30,  1900,  no.  365.  Bemidji,  July  4,  1900,  no.  452, 
July  5,  1900,  no.  483,  July  n,  1900,  nos.  612  and  617  and  July 
14,  1900,  no.  731.  Thief  River  Falls,  July  17,  1900,  no.  768 
and  July  23,  1900,  no.  871.  Red  lake,  July  30,  1900,  no.  998 
and  July  31,  1900,  no.  1008. 

155.  Biatora  rubella  (EHRH.)  RABENH. 

On  elms,  ashes,  poplars,  infrequent.  Battle  lake,  June  20, 
1900,  no.  72.  Henning,  June  25,  1900,  nos.  171,  172  and  173. 
Bemidji,  July  9,  1900,  no.  554,  July  n,  1900,  no.  616,  July  12, 
1900,  no.  681  and  July  13,  1900,  nos.  706  and  712.  Thief 
River  Falls,  July  18,  1900,  no.  783  and  July  21,  1900,  no.  868. 
Red  lake,  July  26,  1900,  no.  892. 

156.  Biatora  fusco-rubella  (HOFFM.)  TUCK. 

On  trees,  infrequent.  Henning,  June  29,  1900,  no.  345.  Red 
lake,  July  28,  1900,  no,  936,  July  31,  1900,  no.  1015  and  Aug. 
i,  1900,  no.  1041. 

157.  Biatora  suffusa  FR. 

On  trees,  rare.     Red  lake,  July  27,  1900,  no.  918. 

158.  Biatora  atrogrisea  (DELIS.)  HEPP. 

On  trees,  infrequent.  Henning,  June  30,  1900,  no.  376. 
Bemidji,  July  n,  1900,  no.  596.  Thief  River  Falls,  July  21, 
1900,  no.  858. 

Not  previously  reported  from  Minnesota. 

159.  Biatora  inundata  FR. 

On  pines  and  rocks,  rare.  Battle  lake,  June  20,  1900,  nos. 
65  and  103.  Bemidji,  July  10,  1900,  no.  575.  Red  lake, 
July  30,  1900,  nos.  981  and  984  and  July  31,  1900,  no.  1040. 

1 60.  Biatora  akompsa  TUCK. 

On  trees,  common.      Battle  lake,  June  18,  1900,  no.  3. 
Not   previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 


Fink:    LICHENS  OF  NORTHWESTERN  MINNESOTA.  703 

161.  Biatora  muscorum  (Sw.)  TUCK. 

On  earth,  frequent.  Battle  lake,  June  19,  1900,  nos.  24 
and  36  and  June  20,  1900,  no.  98.  Leaf  hills,  June  27,  1900, 
no.  285. 

162.  Lecidea  enteroleuca  FR. 

On  trees,  common.  Battle  lake,  June  18,  1900,  no.  7,  June 
20,  1900,  nos.  38  and  61,  June  22,  1900,  no.  130  and  June  23, 
1900,  nos.  151  and  165.  Leaf  hills,  June  27,  1900,  nos.  250 
and  274.  Henning,  June  29,  1900,  no.  341  and  June  30,  1900, 
nos.  363  and  369.  Bemidji,  July  4,  1900,  no.  450,  July  5 ,  1900, 
nos.  466  and  475,  July  n,  1900,  no.  633  and  July  12,  1900, 
nos.  674  and  688.  Thief  River  Falls,  July  17,  1900,  nos.  759, 
764  and  774,  July  19,  1900,  no.  812  and  July  20,  1900,  no. 
843.  Red  lake,  July  27,  1900,  nos.  928  and  932,  July  28, 
1900,  no.  962,  July  31,  1900,  nos.  1004  and  1020  and  August 
i,  1900,  no.  1026. 

163.  Lecidea  enteroleuca  FR.  var.  achrista  SOMMERF. 

On  trees,  infrequent.  Battle  lake,  June  19,  1900,  no.  25. 
Red  lake,  July  28,  1900,  no.  973  and  July  31,  1900,  no.  1009. 

164.  Lecidea  enteroleuca  FR.  var.  flavida  FR. 

On   tamarack   stumps  in   swamps,  rare.     Bemidji,  July  n, 
1900,  no.  610.     Red  lake,  July  28,  1900,  no.  949. 
Not  previously  reported  from  Minnesota. 

165.  Lecidea  enteroleuca  FR.  var.  ambigua  ANZ. 

On  old  wood,  rare.  Battle  lake,  June  23,  1900,  no.  143. 
Henning,  June  25,  1900,  no.  200.  Bemidji,  July  /,  1900,  no. 
518.  Thief  River  Falls,  July  18,  1900,  no.  807. 

Not  previously  reported  from  Minnesota. 

1 66.  Lecidea  acclinis  FLOT. 

On  trees,  infrequent.  Battle  lake,  June  21,  1900,  no.  120. 
Thief  River  Falls,  July  19,  1900,  no.  810  and  July  21,  1900, 
no.  844. 

167.  Lecidea  sp. 

On  dead  tamaracks  in  swamp,  rare.  Henning,  June  25, 
1900,  no.  178.  Thallus  grayish  and  evanescent,  apothecia 
medium  sized ;  hymenium  pale ;  exciple  and  hypothecium 
brownish  black  ;  paraphyses  distinct,  colorless,  filiform,  brown- 
tipped  ;  spores  oblong,  colorless,  4-celled,  straight  or  slightly 
curved,  13-16  mic.  long  by  3^-5  mic.  wide. 


704  MINNESOTA    BOTANICAL    STUDIES. 

168.  Buellia  alboatra  (HOFFM.)  TH.  FR. 

On  trees,  rare.  Battle  lake,  June  21,  1900,  no.  117  and 
June  25,  1900,  no.  158.  Bemidji,  July  u,  1900,  nos.  679  and 
687.  Thief  River  Falls,  July  23,  1900,  no.  872.  Red  lake, 
July  27,  1900,  no.  965. 

169.  Buellia  parasema  (Acn.)  TH.  FR. 

On  trees,  infrequent.  Leaf  hills,  June  27,  1900,  no.  272. 
Bemidji,  July  4,  1900,  no.  434,  July  7,  1900,  nos.  510,  511  and 
542  and  July  12,  1900,  nos.  662  and  696.  Thief  River  Falls, 
July  20,  1900,  no.  831.  Red  lake,  July  28,  1900,  no.  939, 
July  30,  1900,  no.  939  and  August  2,  1900,  no.  1056. 

170.  Buellia  myriocarpa  (DC.)  MUDD. 

On  pines,  rare.  Battle  lake,  June  19,  1900,  no.  23.  Bemidji, 
July  5,  1900,  no.  488.  Red  lake,  August  2,  1900,  no.  1059. 

171.  Buellia  myriocarpa  (DC.)  MUDD.  var.  polyspora  WILLEY. 
On  cedar  in  swamps,  rare.     Bemidji,  July  13,  1900,  no.  714. 

172.  Buellia  pullata  TUCK.  ? 

On  granitic  rocks,  infrequent.  Leaf  hills,  June  26,  1900, 
no.  241. 

Spores  9-14  mic.  long  by  6-7.5  m^c-  wide,  and  thallus 
scanty. 

173.  Buellia  petraea  (FLOT.,  KBR.)  TUCK. 

On  exposed  granitic  rocks,  locally  frequent.  Battle  lake, 
June  23,  1900,  no.  158.  Leaf  hills,  July  2,  1900,  no.  386. 

174.  Buellia   petraea  (FLOT.,   KBR.)   TUCK.    var.   montagnaei 
TUCK. 

On  exposed  granitic  rocks,  locally  common.  Battle  lake, 
June  20,  1900,  no.  62  and  June  23,  1900,  no.  169.  Leaf  hills, 
June  27,  1900,  nos.  283  and  294  and  July  2,  1900,  no.  396. 

175.  Melaspilea  arthonioides  (FEE)  NYL. 

On  trees,  rare.  Battle  lake,  June  21,  1900,  no.  122  and 
June  22,  1900,  no.  133.  Bemidji,  July  n,  1900,  no.  604.  Thief 
River  Falls,  July  21,  1900,  no.  849. 

Not  previonsly  reported  from  Minnesota. 

176.  Opegrapha  varia  (PERS.)  FR. 

On  trees,  common.  Battle  lake,  June  18,  1900,  nos.  15  and 
17  and  June  19,  1900,  nos.  29  and  32.  Henning,  June  25,  1900, 
no.  176  and  June  30,  1900,  no.  362.  Bemidji,  July  4,  1900,  no. 


Fink:   LICHENS  OF  NORTHWESTERN  MINNESOTA.  705 

448.  Thief  River  Falls,  July  18,  1900,  nos.  779  and  790.  Red 
lake,  July  26,  1900,  nos.  881  and  894  and  August  i,  1900, 
no.  1040. 

177.  Graphis  scripta  (L.)  ACH. 

On  trees,  frequent,  Battle  lake,  June  22,  1900,  no.  124. 
Henning,  June  24,  1900,  no.  179.  Bemidji,  July  4,  1900,  no. 
453>  July  JI»  1900*  nos.  603  and  648.  Red  lake,  July  26, 
1900,  no.  880  and  July  30,  1900,  nos.  995  and  997. 

178.  Graphis  scripta  (L.)  ACH.  var.  limitata  ACH. 

On  trees,  rare.  Henning,  June  30,  1900,  no.  367.  Bemidji, 
July  12,  1900,  no.  667. 

179.  Graphis  scripta  (L.)  ACH.  var.  recta  (HuMB.)  NYL. 

On  trees,  rare.  Bemidji,  July  12,  1900,  no.  664  and  July  14, 
1900,  no.  728.  Red  lake,  August  i,  1900,  no.  1037. 

1 80.  Arthonia  lecideella  NYL. 

On  trees,  infrequent.  Battle  lake,  June  18,  1900,  no.  5  and 
June  23,  1900,  no.  161.  Leaf  hills,  June  26,  1900,  no.  242. 
Bemidji,  July  4,  1900,  no.  456,  July  5,  1900,  nos.  463  and  481 
and  July  9,  1900,  no.  552.  Thief  River  Falls,  July  20,  1900, 
no.  837  and  July  2-1,  1900,  no.  856.  Red  lake,  July  31,  1900, 
no.  1013. 

181.  Arthonia  patellulata  NYL. 

On  elms,  rare.     Bemidji,  July  12,  1900,  no.  699. 

182.  Arthonia  dispersa  (SCHRAD.)  NYL. 

On  trees,  probably  common.  Battle  lake,  June  20,  1900, 
nos.  44  and  67,  June  22,  1900,  no.  131  and  June  23,  1900,  no. 
160.  Henning,  June  25,  1900,  no.  183.  Red  lake,  July  26, 
1900,  nos.  895  and  899,  July  28,  1900,  no.  958  and  August  2, 
1900,  no.  1057. 

183.  Arthonia  sp. 

On  trees,  rare.  Leaf  hills,  June  26,  1900,  no.  213.  Spores 
4-celled  in  pyriform  asci,  18-21  mic.  long  by  6-7  mic.  wide. 
Apothecia  differently  disposed  than  in  the  next. 

Not  previously  reported  from  Minnesota. 

184.  Arthonia  radiata  (PERS.)  TH.  FR. 

On  trees,  rare.  Battle  lake,  June  21,  1900,  no.  116  and 
June  22,  1900,  no.  135.  Henning,  June  29,  1900,  no.  360. 
Red  lake,  July  26,  1900,  no.  889  and  July  31,  1900,  nos.  1005 
and  1017. 


706  MINNESOTA    BOTANICAL    STUDIES. 

185.  Acolium  tigillare  (Acn.)  DN. 

An  old  wood,  rare.  Battle  lake,  June  23,  1900,  no.  137. 
Bemidji,  July  5,  1900,  no.  497  and  July  6,  1900,  no.  530. 

1 86.  Calicium  lucidum  (Tn.  FR.)  FINK. 

On  pines,  tamaracks  and  dead  wood,  rare.  Henning,  July 
2,  1900,  no.  413.  Bemidji,  July  6,  1900,  nos.  530  and  539  and 
July  u,  1900,  no.  595.  Red  lake,  July  30,  1900,  no.  994  and 
August  3,  1900,  no.  1072. 

Not  previously  reported  from  Minnesota  and  new  to  North 
America. 

187.  Calicium  trichiale  ACH.  var.  cinereum  NYL. 

On  living  and  dead  wood  in  swamps,  rare.  Henning,  June 
28,  1900,  nos.  301  and  334.  Bemidji,  July  13,  1900,  no.  709. 
Red  lake,  July  26,  1900,  no.  901  and  July  28,  1900,  no.  938. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

188.  Calicium  chrysocephalum  ACH. 

On  cedars  in  swamp,  rare.  Bemidji,  July  6,  1900,  nos.  507 
and  535. 

189.  Calicium  trachelinum  ACH. 

On  old  wood  in  tamarack  swamp,  locally  common.     Henning, 
June  28,  1900,  no.  325  and  July  2,  1900,  no.  406. 
Not  previously  reported  from  Minnesota. 

190.  Calicium  polyporaeum  NYL. 

On  Polyporus  versicolor,  rare.  Bemidji,  July  14,  1900,  no. 
736.  Thief  River  Falls,  July  19,  1900,  no.  823.  Red  lake, 
July  28,  1900,  no.  955. 

Not  previously  reported  from  Minnesota. 

191.  Calicium  sp. 

On  dead  wood,  especially  in  swamps,  rare.  Battle  lake, 
June  23,  1900,  no.  159.  Henning,  June  30,  1900,  no.  374. 
Bemidji,  July  7,  1900,  no.  524  and  July  12,  1900,  no.  692. 

Spores  simple  and  compound.  Referred  to  C.  quercinum 
Pers.  in  no.  IV.  of  this  series  of  papers. 

Not  previously  reported  from  Minnesota. 

192.  Calicium  curtum  TURN,  and  BORR.  ? 

On  old  stumps  of  conifers,  especially  in  swamps,  frequent. 
Henning,  June  28,  1900,  no.  317.  Red  lake,  July  27,  1900, 
no.  911  and  August  2,  1900,  no.  1063. 


Fink:   LICHENS  OF  NORTHWESTERN  MINNESOTA.  707 

Apothecia  are  large  for  the  species  and  not  pruinose  at 
margin. 

Not  previously  reported  from  Minnesota. 

193.  Calicium  trabinellum  (SCHAER.)  KBR. 

On  old  stumps  of  conifers,  common  at  first  locality.  Henning, 
June  28,  1900,  no.  310.  Bemidji,  July  u,  1900,  no.  631. 

Not  previously  reported  from  Minnesota  and  new  to  the  in- 
terior of  North  America. 

194.  Calicium  parietinum  ACH. 

On  dead  stumps  of  conifers,  common  especially  in  swamps. 
Henning,  June  28,  1900,  nos.  299  and  316.  Bemidji,  July  7, 
1900,  no.  523  and  July  12,  1900,  no.  678.  Thief  River  Falls, 
July  21,  1900,  no.  866.  Red  lake,  July  27,  1900,  no.  916  and 
August  i,  1900,  no.  1035. 

195.  Calicium  pusillum  FLK. 

On  dead  tamarack  in  swamps,  rare.  Henning,  June  25, 
1900,  no.  196. 

Not  previously  reported  from  Minnesota  and  new  to  North 
America. 

196.  Calicium  turbinatum  PERS. 

On  Pertusaria  communis,  rare.     Bemidji,  July  13,  1900,  no. 

7°3- 

197.  Coniocybe  pallida  (PERS.)  FR. 

On  elms,  infrequent.  Henning,  June  30,  1900,  no.  368. 
Bemidji,  July  u,  1900,  no.  630  and  July  13,  no.  708.  Thief 
River  Falls,  July  19,  1900,  no.  716.  Red  lake,  July  31,  1900, 
no.  1007. 

198.  Endocarpon  fluviatile  DC. 

On  rocks  by  water,  infrequent.  Red  lake,  August  2,  1900, 
no.  1052. 

199.  Endocarpon  hepaticum  ACH. 

On  earth,  usually  calcareous,  abundant  at  second  locality. 
Battle  lake,  June  20,  1900,  no.  95.  Leaf  hills,  June  27,  1900, 
nos.  256  and  273  and  July  2,  1900,  nos.  389  and  398.  Bemidji, 
July  16,  1900,  no.  742. 

200.  Endocarpon  pusillum  HEDW. 

On  calcareous  pebbles,  locally  frequent.  Leaf  hills,  June 
27,  1900,  no.  258  and  July  2,  1900,  no.  398. 


708  MINNESOTA    BOTANICAL    STUDIES. 

201.  Staurotheleumbrina  WAHL.)TUCK.  var.  colpima(WHNBL.), 

NYL. 

On  granitic  rocks  in  damp  places  and  usually  near  water, 
locally  common.  Bemidji,  July  n,  1900,  nos.  593,  636,  641 
and  642. 

This  plant  was  recorded  in  no.  IV.  of  this  series  of  papers 
and  also  in  no.  V.  as  S.  drummondii  Tuck.  However,  it 
agrees  more  nearly  with  European  specimens  of  the  present 
variety. 

Not  previously  reported  from  Minnesota. 

202.  Verrucaria  nigrescens  PERS. 

On  rocks,  frequent.  Battle  lake,  June  20,  1900,  nos.  86 
and  87  and  June  23,  1900,  nos.  144  and  168.  Leaf  hills,  July 
2,  1900,  nos.  387  and  403.  Thief  River  Falls,  July  17,  1900. 
no.  754.  Red  lake,  July  27,  1900,  no.  929  and  July  28,  1900, 
no.  947. 

203.  Verrucaria  viridula  ACH. 

On  rocks  along  lake,  common.  Red  lake,  July  28,  1900, 
no.  943. 

Not  previously  reported  from  Minnesota. 

204.  Verrucaria  muralis  ACH. 

On  rocks,  especially  calcareous,  infrequent.  Battle  lake, 
June  23,  1900,  no.  166.  Leaf  hills,  June  27,  1900,  nos.  289 
and  293  and  July  2,  1900,  no.  404.  Red  lake,  July  28,  1900, 
no.  940. 

205 .  Verrucaria  fuscella  FR. 

On  calcareous  drift  pebbles,  frequent.     Leaf  hills,  June  27, 
1900,  nos.  262  and  297. 
A  peculiar  form. 

206.  Verrucaria  conoidea  FR. 

On  lime  pebbles,  rare.     Bemidji,  July  n,  1900,  no.  626. 
Not  previously  reported  from  Minnesota  and  probably  new  to 
North  America. 

207.  Lagedia  oxyspora  (NYL.)  TUCK. 

On  birch  trees,  probably  abundant.  Henning,  June  25,  1900, 
no.  189.  Bemidji,  July  n,  1900,  no.  608  and  July  12,  1900, 
no.  671. 

208.  Pyrenula  punctiformis  (Acn.)  NAEG. 

On  trees,  rare.     Battle  lake,  June  18,  1900,  no.  18. 


Fink:   LICHENS  OF  NORTHWESTERN  MINNESOTA.  709 

209.  Pyrenula  punctiformis  (Acn.)  NAEG.  var.  fallax  NYL. 

On  birch  trees,  common.  Henning,  June  25,  1900,  no.  170 
and  June  29,  1900,  no.  353.  Bemidji,  July  n,  1900,  nos.  601 
and  624  and  July  12,  1900,  no.  670.  Red  lake,  August  i,  1900, 
no.  1036. 

The  more  I  see  of  this  plant,  the  more  likely  it  seems  that  it 
will  finally  have  to  be  separated  entirely  from  this  species. 

210.  Pyrenula  gemmata  (AcH.)  NAEG. 

On  trees,  rare.     Thief  River  Falls,  July  18,  1900,  no.  798. 

211.  Pyrenula  hyalospora  (NYL.)  TUCK. 

On  trees,  locally  frequent.  Red  lake,  July  26,  1900,  no. 
887. 

212.  Pyrenula  cinerella  (FLOT.)  TUCK. 

On  birch,  locally  common.     Henning,  June    29,   1900,  no. 

348. 

213.  Pyrenula  cinerella    (FLOT.)    TUCK.    var.    quadriloculata 
FINK. 

On  birch,  common.  Bemidji,  July  n,  1900,  nos.  600  and 
632.  Red  lake,  July  26,  1900,  no.  897. 

214.  Pyrenula  leucoplaca  (WAHL.)  KBR. 

On  trees,  especially  poplars,  common.  Leaf  hills,  June  27, 
1900,  nos.  268  and  269.  Henning,  June  29,  1900,  no.  344. 
Bemidji,  July  9,  1900,  no.  555,  July  u,  1900,  no.  611  and  July 
12,  1900,  no.  676.  Thief  River  Falls,  July  17,  1900,  no.  767, 
July  18,  1900,  nos.  789  and  802  and  July  19,  1900,  no.  821. 
Red  lake,  July  28,  1900,  no.  961. 

215.  Pyrenula  leucoplaca  (WAHL.)  KBR.  var.  pluriloculata  var. 
nov. 

On  trees,  infrequent.  Battle  lake,  June  18,  1900,  no.  10 
and  June  21,  1900,  no.  in.  Henning,  June  29,  1900,  no. 
357.  Bemidji,  July  u,  1900,  no.  597.  Thief  River  Falls, 
July  18,  1900,  no.  799  and  July  19,  1900,  no.  829.  Red  lake, 
August  3,  1900,  no.  1073. 

Spores  5-8-celled. 


XL.  CORALLINE  VER^  OF  PORT  RENFREW. 


K.  YENDO,  Rigakushi, 
SCIENCE  COLLEGE,  IMPERIAL  UNIVERSITY,  TOKYO. 


I  had  the  opportunity  of  studying  seaweeds  during  the  sum- 
mer of  1901,  at  the  Minnesota  Seaside  Station  at  Port  Ren- 
frew, Vancouver  Island,  B.  C.  The  vicinity  of  the  laboratory 
is  extremely  rich  in  marine  life  and  afforded  a  good  number 
of  examples.  I  paid  special  attention  to  calcareous  algae,  in 
which  branch  I  have  been  deeply  interested.  Returning  with 
the  specimens  to  the  Botanical  Institute  of  the  University  of 
Tokyo,  I  carefully  examined  the  Corallines  (verae)  and  pre- 
pared the  present  paper.  Other  specimens  belonging  to  the 
subfamily  Melobesiae  have  been  sent  to  Mr.  M.  Foslie,  of  Nor- 
way ;  his  paper  should  also  appear  in  a  short  time. 

The  material  was  partly  dried  and  partly  preserved  in  alco- 
hol, the  accompanying  photographs  being  taken  from  the  latter. 
The  sections  were  made  from  alcoholic  material  by  microtome, 
after  decalcifying  in  Pereny's  fluid.  Amphfroa  tuberculosa  and 
a  few  other  thick  plants  were  not  satisfactorily  decalcified  by 
the  solution  and  the  author  found  the  following  mixture  specially 
suited  for  the  purpose  : 

Hydrochloric  acid,  5   per  cent 40  c.cm. 

Alcohol,  absolute 30  c.cm. 

Chromic   acid,  0.5   per  cent 30  c.cm. 

The  sections  were  stained  after  my  usual  method.  One 
brings  down  the  sections  to  pure  water,  and  stains  with  Boemer's 
haematoxylin  for  20—40  minutes  ;  treat  with  acetic  acid  if  neces- 
sary, and  then  dip  in  fuchsin  (0.3  gr.  in  100  c.cm.  of  50  per 
cent,  alcohol  for  one  hour;  90  per  cent,  alcohol,  abs.  alcohol, 
xylol,  balsam.  The  spores  and  spore-forming  cells  stain  in  red 
and  the  cell-walls  of  the  vegetative  cells  in  purple. 

711 


712  MINNESOTA    BOTANICAL    STUDIES. 

The  author  expresses  his  deepest  thanks  to  Professor  Conway 
MacMillan  and  Miss  J.  E.  Tilden,  who  cared  for  him  very 
kindly  in  every  way  during  his  stay  at  their  private  station  ;  and 
to  Professor  J.  Matsumura,  of  Tokyo,  who  offered  many  valu- 
able suggestions  on  this  work. 

SYNOPTICAL  KEY  OF  CORALLINE  ENUMERATED  IN  THE 
PRESENT  PAPER. 

A.  Conceptacles  wart-like  protuberances,  on  a  surface  of  an  articulus. 

o.   Branches  not  pinnated. 

a.  Articuli  cylindrical Amphiroa  cretacea  L  tasmanica . 

b.  Articuli  of  the  upper  and  the  middle  portions  cylindri- 
cal, subcompressed  or  compressed. 

Amphiroa  tuberculosa. 

c.  Articuli  of  the  upper  and  middle  portions  approximate, 
with  depressed  wings Cheilosporum  californicum. 

00.   Branches  pinnated. 

a.  With    short    and    thick    stipes;     upper    articuli    of    the 
branches  cordate  or  sagittate  with  round  lobes;  external 
margins  of  the  ultimate  articuli  thick. 

Cheilosporum  frondescen  <•. 

b.  With  long  and  thin  stipes ;  upper  articuli  of  the  branches 
sagittate  with  acute  lobes ;   external  margins  of  the  ulti- 
mate articuli  thin Cheilosporum  planiusculum. 

c.  With  long  and  thick  stipes ;  articuli  of  the  axial  branches 
hexagonal,  without  evident  rib ;   conceptacles  sometimes 
stalked Cheilosporum  MacMillani 

B.  Conceptacles  stalked,  taking  place  of  segments. 

o.  Branches  regularly  pinnated,  flabellate. 

Corallina  ojficinalis  var.  chilensis. 

oo.  Branches    irregularly  pinnated,   several  pinnules  around  the 
top  of  an  articulus. 

a.  Pinnules  not  confusedly  ramous. 

Corallina  vancouveriensis. 

b.  Pinnules  confusedly  ramous  and  prickly. 

Corallina  aculeata. 

HABITAT  OF  CORALLINE  AT  PORT  RENFREW. 

The  coast  near  the  Minnesota  Seaside  Station  chiefly  consists 
of  sandstone  beds  spreading  horizontally.  The  beds  are  cov- 
ered with  water  at  high  tide,  and  drained  at  low  tide,  leaving 
a  good  number  of  pools.  The  Corallinae  are  mostly  found 


Tendo :  CORALLINAE  VER^E.  713 

between  the  tidal  marks  as  well  as  in  the  pools.  Amphiroa 
tuberculosa  and  Amp.  cretacea  f.  tasmanica  are  found  at  the 
depth  of  2-5  ft.  below  the  low-water  mark  or  the  surface  of  the 
pools  :  the  former  species  assumes  very  diverse  forms  when  it 
is  found  at  the  margins  of  the  pools,  or  between  tidal  marks. 
So  also  do  Corallina  officinalis  var.  chilensis,  and  Cheilosporum 
MacMillani ;  but  the  latter  two  are  not  infrequently  found  in 
more  shallow  water.  Cor.  vancouveriensis  and  its  variety,  on 
the  contrary,  are  in  most  cases  found  at  the  margins  of  the 
pools,  and  in  the  region  a  little  above  the  low-tide  mark.  They 
are  also  found  epiphytic  on  the  shells  of  Mya  which  cover  the 
enormous  area  of  the  drained  beds,  thus  making  it  easy  to  col- 
lect the  entire  bunch  of  the  plants.  Cheil.  frondescens  is  also 
found  in  similar  positions.  ChciL  planiusculum  is  also  an  in- 
habitant of  the  pools,  but  slightly  below  the  margins.  When 
it  grew  above  the  water  mark  the  frond  is  mostly  stunted,  appar- 
ently forming  a  granular  mass.  In  the  tide  pools  high  above 
the  water-level  Cor.  aculeata  is  generally  found  ;  water  in  such 
pools  is  mostly  brackish,  at  least  during  the  ebb  tide  hours  ;  and 
the  plant  seemed  to  be  able  to  adapt  itself  to  it.  This  might  be 
the  probable  cause  why  the  ultimate  articuli  of  the  branches  of 
this  species  are  insufficiently  calcified.  Nevertheless,  Cor.  van- 
couveriensis,  Cheil.  calif ornicum,  Cheil.  planiusculum,  etc.,  may 
also  be  found  in  these  brackish  pools  without  any  apparent  modi- 
fication in  the  characters  of  their  fronds.  Briefly  speaking,  Cor. 
vancouveriensis  is  an  inhabitant  of  the  shallowest  water,  and 
Amp.  tuberculosa^  as  it  were,  of  the  deepest.  The  latter  view 
may  be  corroborated  by  the  fact  that  we  often  find  the  frag- 
ments of  Amp.  tuberculosa  growing  attached  to  the  holdfast  of 
Ncreocystis  Lutkeana  hauled  up  out  of  water  20-50  feet  deep. 

Cor.  pilulifera  and  its  varieties  which  are  abundantly  found 
in  Kamtchatka  and  in  the  northern  part  of  Japan  could  not  be 
found  at  Port  Renfrew.  Their  places  seem  to  be  taken  by 
Cor.  vancouveriensis  and  Cheil.  planiusculum.  The  habitat 
of  these  is  much  like  that  of  the  typical  form  of  Cor.  officinalis 
or  Cor.  squamata. 

Cheil.  frondescens  which  was  described  by  Ruprecht*  col- 
lected in  Unalaska  is  common  at  this  coast.  Areschoug  f  re- 


*Post.  et  Rupr.:  Illustr.  alg.,  p.  20. 

t  Aresch.:  in  J.  Ag.  Spec,  alg.,  II.,  p.  549. 


714  MINNESOTA    BOTANICAL    STUDIES. 

marked  Ruprecht's  plant  to  be  probably  a  dwarfed  form.    This 
.  remark  is  true  so  far  as  my  determination  is  correct ;   our  plant 
is  little  larger  than  Ruprecht's  f .  maxima  mihi  being  twice  or 
more  as  large  as  his  plant. 

No  specimens  of  Jania  could  be  found.  The  water  at  Port 
Renfrew  seems  to  be  too  cold  to  admit  any  member  of  this 
group.  We  are  able  to  give  only  one  datum  here  :  the  tem- 
perature of  the  water  y2  ft.  below  the  surface  of  a  tidal  pool, 
io°.4  C.  ;  i  ft.  below  the  surface  of  open  sea  io°.2  C.  ;  Atm. 
temp.  u°. 55  C. 

CORALLINE  VER^E  OF  PORT  RENFREW. 

1.  Amphiroa  cretacea  Endl.  f.tasmanica(Sond.).    PI. LI., Fig.  i. 
=  Amphiroa  tasmanica  Sond.  in  Plant  Mull.  (Linnsea,  XXV.). 

2d  :  in  Mull.  Frag.  Phyt.  Austr.  Suppl. 
Kiitzing:  Tab.  Phyc.,  VIII.  Taf.  47,  Fig.  n. 
The  plant  found  at  Port  Renfrew  is  identical  with  the  Tas- 
manian  form  and  not  with  Amp.  cretacea,  which  was  collected 
in  Unalaska  by  Ruprecht.     As  has  been  already  remarked  by 
Kiitzing  (/.  c.,  p.  23),  Amp.  tasmanica  Sond.  is  quite  similar  to 
Amp.  cretacea  Endl.  and  it  might  better  be  reduced  as  above. 
Not  rare  :   2—5  ft.  below  low-water  mark,  also  in  pools. 

2.  Amphiroa  tuberculosa  Endl.    PL  LI.,  Fig.  2  ;  PL  LVL,  Figs. 

i  and  2. 

Aresch  :  in  J.  Ag.  Spec.  Alg.,  II.,  p.  538. 
Harv :  Ner.  Bor.  Am.,  p.  86. 

=  Corallina  tuberculosa  Post,  et  Rupr.  111.,  p.  20,  t.  40. 
Kiitz  :  spec,  alg.,  p.  704. 

?  =  Amphiroa  (Artkrocardid]  cpiphlegnoides  J.  Ag.  in  Har- 
vey's Notes  on  N.W.  Am.  Alg.Qourn.of  Linn.  Soc.,VI.,p.  169). 

=  Amphiroa  calif ornica  in  Prov.  Museum  at  Victoria,  B.  C. 

Judging  by  the  figure  delineated  by  Postels  and  Ruprecht 
(/.  c.}  our  plant  may  be  readily  referred  to  the  present  species. 
It  attains  to  3-5  inches  in  its  height  with  subdichotomous  or 
lateral  patent  branches.  The  articuli  are  extremely  variable  in 
their  form  :  those  of  the  basal  portion  are  invariably  subcylin- 
drical ;  those  of  the  upper  and  the  middle  portions,  cordati  or 
sagittate,  sometimes  cylindrical  or  clavate  ;  the  cordate  or  sagit- 
tate articuli  are  more  or  less  compressed  and  generally  \\ith 
subevident  rib  on  the  shaded  surface  :  the  terminal  articuli  are 


Yendo :    CORALLINE  VER^E.  715 

normally  subcompressed  obovate  but  sometimes  globular  or 
linear.  The  conceptacles  are  slightly  bulged  out,  two  to  several, 
mostly  immersed  in  the  shaded  side  of  a  cordate  articulus. 
Geniculi  lineaeform. 

A  branch  is  often  simple  with  homogeneous  cylindrical  ar- 
ticuli.  Plants  with  lots  of  this  sort  of  branch  are  likely  to  be 
confounded  with  Amp.  cretacea  Endl.  or  the  preceding  forma. 
But  the  occurrence  of  the  cordate  articuli  is  the  character  upon 
which  to  separate  the  present  species.  Amp.  epiphlegnoides 
set- ms  to  me  quite  similar  to  this  plant.  I  mention  it  here  as  a 
synonym,  however,  with  doubt,  as  I  have  not  seen  any  authentic 
specimen  of  Agardh's  plant. 

Common  :  2—5  ft.  below  low  water  mark,  not  seldom  found, 
several  fathoms  deep. 

3.  Cheilosporum  californicum  (Dene.).     PL  LIV.,  Fig.  2  ;  PI. 
LVI.,  Fig.  3. 

Frond  3-5  cm.  alta,  stipitata,  irregulariter  di-trichotome  ra- 
mosa :  articulis  stipitis  cylindraceis  diametro  sesqui-  2-  plo 
lorigioribus  sursum  latioribus  et  compressis ;  ramorum  approxi- 
matis,  adpressis  mediis  costatis,  obreniformibus  vel  sagittatis 
lobis  rotundis,  longitudine  inter  genicula  distantiam  loborum 
4-plo  brevioribus  ;  apicalium  obovatis  compressis  :  conceptaculis 
hemisphericalibus  binis  vel  quatuor  in  utraque  facie  articuli 
instructis. 

Amphiroa  californica  Dene.,  Class  f.  d.  Alg.  et  Cov.,  p.  112. 
Kiitz  :  Spec.  Alg.,  p.  704. 
Aresch  :  in  J.  Ag.  Spec.  Alg.,  II.,  p.  542. 
Harv:   Ner.  Bor.  Americ,  p.  86. 

As  the  original  description  of  this  plant  was  given  somewhat 
briefly,  a  few  other  species  have  been  mistakenly  identified  with 
it;  and  Areschoug  counted  it  under  the  "species  inquirendae" 
(/.  c.).  We  have  a  specimen  of  Amp.  californica  Dene.,  col- 
lected at  Oregon  and  determined  by  Dr.  Farlow.  Our  plant  is 
exactly  similar  to  this  specimen  and  at  once  accords  very  well 
with  Decaisne's  description. 

Not  rare  :   low-water  mark,  also  in  tide  pools. 

4.  Cheilosporum  frondescens  (Post,  et  Rupr.),  f.  typica.     PI. 
LIL,  Fig.  i  ;  PI.  LVI.,  Figs.  4,  5  and  8. 

C or  allina  frondescens  Post,  et  Rupr.,  111.  p.  20,  t.  XL.,  f.  103. 


716  MINNESOTA    BOTANICAL    STUDIES. 

Arthrocardia  f  frondesccns  (Post,  et  Rupr.)  Aresch.  in  J.  Ag. 
Spec.  Alg.,  II.,  p.  549. 

f.  maxima,  f.  nov.     PI.  LIL,  Fig.  3. 

Fronde  majore  et  crassiore  ;  articulis  pinnarum  lobis  latissimis 
saepe  crenulatis,  apicalium  compressis  obovatis  vel  spatulatis. 

f.  intermedia,  f.  nov.     PI.  LIL,  Fig.  2. 

Fronde  tenuiore ;  articulis  pinnarum  compressis  deltoideo- 
obcordatis  lobis  nonnunquam  acutis. 

f.  polymorpha,  f .  nov.  PI.  LIL,  Fig.  4;  PI.  LVL,  Figs.  6, 
6a  and  7. 

Fronde  minore  ;  crassiuscula,  polymorpha  :  turn  articulis  pin- 
narum obcordatis,  obreniformibus  vel  sagittatis,  apicalium  obo- 
vatis vel  globosis ;  turn  articulis  pinnarum  axiumque  cylindra- 
ceis,  linearibus  vel  globosis. 

Ruprecht's  illustrations  and  description  /.  c.  precisely  coincide 
with  f.  typica.  Areschoug  remarked  in  J.  Ag.  Spec.  Alg.  /.  c. 
that  the  original  plant  might  have  been  a  tiny  form  of  the  spe- 
cies. We  have  at  Port  Renfrew  plants  often  attaining  to  several 
centimeters  in  height. 

f .  typica  is  a  densely  caespitose  plant,  irregularly  pinnated ; 
the  lowermost  pinnae  attain  to  the  length  of  the  main  stem,  and 
thus  give  the  appearance  of  trichotomy.  The  articuli  are  com- 
pactly arranged,  the  lower  margin  of  an  articulus  in  contact 
with  the  upper  margin  of  the  subsequent  one.  Conceptacles 
are  mostly  found  two  to  four  in  number  and  placed  on  the 
shaded  surface,  often,  however,  solitarily  immersed  at  the 
angles  of  a  deltoid  articulus. 

f.  maxima  is  distinguished  from  the  other  formae  by  its  large 
and  compressed  articuli  at  the  upper  portion  of  the  branches. 
These  articuli  measure  2-5  mm.  broad,  1.5-2  mm.  long  and 
are  often  cleft  at  their  lobes.  The  branches  are  not  so  dense  as 
in  f .  typica. 

f.  intermedia  is  characterized  by  having  the  upper  portions  of 
the  fronds  revoluted  downwards  while  it  is  yet  young.  The 
articuli  are  rather  less  wide  than  those  of  f.  typica  and  in  every 
part  thin.  The  lobes  of  the  articuli  are  angled  and  consequently 
more  loosely  arranged. 

f .  polymorpha  attains  to  a  length  of  scarcely  one  inch  and 
has  its  articuli  thick  and  rough.  Its  form  is  very  variable,  some- 
times assuming  quite  an  aberrant  appearance  (PL  LVL,  Fig.  6, 


Tendo:    CORALLINE  VERJE.  717 

6a)  so  that  we  could  not  suppose  it  to  belong  to  this  species, 
had  it  not  been  provided  with  some  normal  branches  in  a  portion 
of  the  frond  (PL  LIL,  Fig.  4). 

Although  I  distinguish  these  four  formae,  intermediate  forms 
between  them  are  naturally  met  with.  Especially  f.  intermedia 
and  f .  -polymorpha  are  likely  to  be  confounded  with  the  abnormal 
forms  of  Cheil.  planiusculum.  In  this  case  the  external  thick 
margin  of  the  apical  articuli  and  the  robust  stipes  are  the  impor- 
tant characters  of  this  species  to  separate  it  from  the  latter.  The 
apical  articuli  of  Chcil.  -planiusculum  are  mostly  thin  and  com- 
pressed, and  the  stipes  are  delicate  filiform.  Nevertheless,  it 
would  not  be  an  unreasonable  supposition  that  the  hybrid  between 
Chcil.  frondcsccns  and  Chcil.  planiusculum  ma)7  occur  in  nature. 

Common :    between  tide  marks,  also  in  pools. 

5.  Cheilosporum  planiusculum  (Kiitz.).    PL  LIIL,  Figs.   1-3; 

PL  LVI.,  Figs.  9  and  10. 

Fronde  dense  ca^spitosa  suberecta,  3-7  cm.  alta,  superne  com- 
planata,  bi-tripinnata  ;  articulis  axium  inferioribus  tenuioribus 
cylindraceis  mediis  superioribusque  compressis  late  triangulari- 
bus  subcostatis,  pinnarum  sagittatis  lobis  acutis  saspe  cordatis, 
pinnularum  ancipitibus  lanceolatis  vel  linearibus,  ultimis  obo- 
vatis  compressis ;  geniculus  brevissimis ;  conceptaculis  hemi- 
sphaericalibus,  2-5  in  articulo  instructio. 

Corallina  planiuscula  Kiitz.,  Tab.  Phyc.,  VIII.,  p.  3i,taf. 
63>  Fig.  3. 

The  present  plant  is  extremely  variable  in  the  shape  of  its 
articuli,  and  sharp  definition  is  hard  to  give.  Kiitzing  counted 
four  formae  in  the  original  description  (/.  c.)  though  I  could  not 
find  any  form  referable  to  f.  laciniata.  The  other  three  formae 
may  be  found  mixed  together  in  one  bunch  of  the  plant,  often 
branches  of  different  forms  occurring  in  one  individual.  In  an 
extreme  instance,  especially  in  a  plant  growing  at  high-tide 
mark,  the  frond  becomes  a  moniliform  filament  with  a  few 
articuli  of  the  normal  shape  (Plate  LIL,  Fig.  3).  Generally 
speaking,  the  articuli  of  the  upper  and  middle  portions  are 
sagittate,  with  lobes  thin,  delicate  and  sharp  at  the  upper  angles, 
and  with  evident  ribs  at  the  middle ;  the  pinnules  are  thin, 
spatulate  or  lanceolate.  As  the  consequence,  an  articulus  is 
not  approximate  with  its  adjacent  ones  as  in  Cheil.  frondescens 
f.  typtca(comp.  PL  LVI.,  Fig.  4,  and  PL  LVI.,  Fig.  10). 


718  MINNESOTA    BOTANICAL    STUDIES. 

The  delicate  moniliform  stipes,  and  the  thin  external  margin 
of  the  apical  articuli  are  the  important  characters  of  this  spe- 
cies to  distinguish  it  from  the  preceding  species. 

Comparatively  common  ;  low  tide  mark,  also  in  pools. 

6.  Cneilosporum  MacMillani  sp.  nov.  Plate  LII.,  Figs.  4  and  5  ; 
PI.  LVL,  Figs.  11-14. 

Fronde  crassa  nudusculo-stipitata,  4-10  cm.  alta,  inferne  sub- 
teretibus  superne  flabellata  bi-tripinnata ;  pinnis  pinnulisque 
creberrimis  erecto-patentibus  sursum  sensim  brevioribus  :  articu- 
lis  stipitum  cylindraceis  diametrum  subaequantibus,  mediis  et 
superioribus  compressis  medio  ventro  elevatis,  hexagonis  vel 
cuneato-deltoideis  diametro  sesquibrevioribus,  pinnarum  anci- 
pitibus  lineari-sagittatis  vel  lanceolatis,  ultimio  ellipsoideis 
saepe  incrassatis ;  conceptaculis  verrucgeformibus  in  utraque 
facie  instructis  vel  subcompressis  in  apicibus  pinnularum  im- 
mersis,  nonnunquam  pyriformibus  pedunculatis. 

This  plant  has  its  articuli  of  the  upper  part  of  the  main 
branches  more  highly  elevated  on  the  ventral  side  than  on  the 
dorsal.  The  articuli  of  the  lower  portion  are  thick  and  monili- 
form, gently  compressed  upwards :  in  the  upper  and  middle 
portion  they  become  hexagonal  or  truncated,  shorter  than  the 
breadth. 

According  to  the  description*  of  Amphiroa  ivardii  Harv.  and 
Amp.  mallardice  Harv.,  the  present  species  has  some  common 
characters  with  them  so  that  I  hesitated  for  a  time  to  name  it  as 
a  new  species.  But  the  attachment  of  the  conceptacles  of  our 
plant  is  rather  peculiar,  only  one  similar  example  being  hitherto 
known  in  Cheil.  maximum.^  The  eramiferous  articuli  of  the 
pinnae  of  ours  are  lineari-sagittate  with  lobes  projected  upwards  ; 
and  the  articuli  of  the  upper  portions  of  the  main  branches  are 
hexagonal,  much  broader  than  the  height.  These  various  char- 
acters may  easily  distinguish  ours  from  Harvey's  plants.  Not 
common  ;  low-tide  mark. 

7.  Corallina  officinalis  var.  chilensis  Kutz.     PI.  LIV.,Fig.  i; 
PI.  LVL,  Fig.  15. 

Fronde  erecta,  5-10  cm.  alta,  inferne  teretiuscula,  superne 
flabeliata  bi-tripinnata :  articulis  inferioribus  compressiusculis, 


*  Harvey  :  Nereis  Austr.,  p.  99. 

|Yendo:  Cor.  verae  Japan.  (Journ.  ofSci.  Coll.  Tokyo,  vol.  XVI.,  art.  III.) 


Tendo:    CORALLINE  VEUVE.  719 

mediis  superioribusque  oblongo-cuneatis  compressis,  pinnarum 
sterilium  linearibus  vel  lanceolatis  ancipitibus,  ultimis  compres- 
sis obovatis ;  conceptaculio  pedunculatis  subcompressis  saepe 
corniculatis.  Color  rubro-violaceus. 

Corallina  officinalis  chilensis  Kiitz.,  Tab.  phyc.,  VIII.,  p.  32, 
taf.  66,  Fig.  i. 

Cor.  officinalis  L.  f.  d  Yendo.  Cor.  verae  Japan.,  PI.  VII., 
Fig.  13  (Journ.  Sc.  Coll.  Tokyo,  Vol.  XVI.). 

The  sterile  specimens  of  this  variety  have  been  collected  at 
Hakodate,  a  port  in  the  northern  part  of  Japan.  As  they  lacked 
the  conceptacle  I  was  not  able  to  satisfactorily  determine  the 
species  and  included  them  under  the  Cor.  officinalis  L.  The 
specimens  collected  at  Port  Renfrew  are  fortunately  fertile  and 
accord  very  well  with  the  description  and  figures  of  Kiitzing's 
Tab.  Phyc.  and  at  the  same  time  correspond  with  the  Hakodate 
specimens. 

As  I  before  noted  (/.  c.),  this  plant  is  a  somewhat  variable 
form  to  be  counted  under  the  species  Cor.  officinalis  L. 

Not  very  common  ;  low-water  mark,  also  in  pools  at  the  depth 
of  2-3  ft.  below  the  surface. 

8.  Corallina  vancouveriensis  sp.  nov.     PI.  LIV.,  Fig.  3  ;  PI. 

LV.,  Figs,  i  and  2  ;  PL  LVI.,  Figs.  16-17. 

Fronde  5-15  cm.  alta,  multicipite,  longe  stipitata,  ramis  bi-tri- 
pinnatis,  ssepe  pinnulis  ex  apice  articuli  egredientibus  ;  articulis 
infinis  globosis,  mediis  superioribusque  subclavatis  diametro 
aequalibus  vel  2-plo  longioribus  tereti-compressis,  ultimis  obo- 
vatis subcompressis  ;  articulis  pinnarum  cylindraceis  linearibus 
vel  alato-projectis  digitalis  ;  conceptaculis  globosis  vel  pyriform- 
ibus  stipitatis,  saepe  corniculatis. 

f.  typica,  f.  nov.     (PI.  LIV.,  Fig.  3  ;  PL  LVI.,  Fig.  16.) 

Fronde  plena  articulorum  linearum  vel  alato-projectorum  digi- 
tatorum,  conceptaculis  globosis  longe  stipitatis. 

f.  densa,  f.  nov.     (PL  LV.,  Fig.  i ;  PL  LVI.,  Fig.  17.) 

Fronde  dense  ramosa, conceptaculis  pyriformibus  pedunculatis. 
Both  formae  approach  one  another  and  a  sharp  boundary  is 
hard  to  draw.  But  f.  typica  is  thicker  and  larger  than  the 
other  and  has  abundance  of  linear  or  lanceolate  long  pinnae  in 
the  upper  part  of  the  frond.  The  high  tide  form  of  this  species 
assumes  a  diverse  appearance ;  its  stipes  are  thick  and  stunted, 


720  MINNESOTA    BOTANICAL    STUDIES. 

the  pinnules  in  the  upper  portion  are  robust  and  fan-shaped, 
generally  crenulated  at  the  external  margin. 

Most  common  between  tide  marks  on  the  margins  of  the  pools. 

9.    Corallina  aculeata  sp.  nov.     PL  LV.,  Fig.  3;    PL  LVL, 

Figs.  18-19. 

Fronde  5-10  cm.  alta,  stipitata,  irregulariter  bi-tripinnata, 
saepe  pinnulis  ex  apice  articuli  egredientibus ;  articulis  inferiori- 
bus  diametro  sesquilongioribus,  pinnarum  pinnularumque  fragil- 
issimis  digitato-laciniatis  aculeatio,  ssepe  cylindraceis  vel  line- 
aribus ;  conceptaculis  subcompressis  cornibus  aculeatis. 

The  pinnules  of  this  plant  are  characteristic  :  they  are  brittle, 
delicate  and  confusedly  branched.  The  ultimate  articuli  of  the 
main  branches  as  well  as  some  of  the  young  pinnules  are  al- 
ways weakly  calcified ;  and  the  apices  of  these  articuli  are 
liable  to  shrink  in  the  exsiccation.  In  other  respects  it  is 
closely  allied  to  Cor.  vancouveriensis  f.  typt'ca^  so  that  it  might 
be  taken  as  a  local  form  caused  by  the  mode  of  habit.  Indeed 
a  young  and  sterile  frond  of  this  species  is  hardly  separable  from 
it,  if  the  apical  articuli  were  not  weakly  calcified. 

Common;  high-tide  pools. 

EXPLANATION  OF  PLATES. 

The  figures  in  Plates  LI.-LV.  are  all  in  natural  size,  taken  from 
the  alcoholic  specimens. 

PLATE  LI. 

1.  Amphiroa  cretacea  Endl.  var.  tasmanica  Sond. 

2.  Amphiroa  tuberculosa  Endl. 

PLATE  LII. 

1.  Cheilosporum frondescens  (Post,  et  Rupr. )  f.  typica. 

2.  do.  f.  intermedia. 

3.  do.  f.  maxima. 

4.  do.  f.  polymorpha. 

PLATE  LIII. 

1.  Cheilosporum  planiusculum  (Kiitz.)  f.  regularis. 

2.  do.  f.  normalis. 

3.  An  abnormal  form  of  the  same  species  found  at  high-tide  mark. 
The  fronds  are  filamentous  with  moniliform  articuli;   a  few  sagittate 
articuli  are  to  be  found  in  some  parts. 


Yendo:   CORALLINE:  VER.K.  721 

4  and  5.  Cheilosporum  MacAIillani.  In  figure  4  wart-like  con- 
ceptacles  as  well  as  pyriform  ones  are  to  be  found  on  the  same  branch. 

PLATE  LIV. 

1.  Corallina  officinalis  L.  var.  chilensis  Kiitz 

2.  Cheilosporum  calif ornicum  (Dene.). 

3.  Corallina  Vancouver lens is  f.  typica. 

PLATE  LV. 

1.  Corallina  vancouveriensis  f.  densa. 

2.  A  high-tide  form  of    Corallina  Vancouver  iensis. 

3.  Corallina  aculeata. 

PLATE  LVI. 
Figures  i  and  2.      Amphiroa  tuberculosa  Endl. 

1.  Diagrammatic  figure  of  the  longitudinal   section  of  an  articulus 
showing  four  conceptacles,  three  of  which  are  cut  in  meridional  direc- 
tion, the  remaining  one  in  crosswise,      x  ca.  15. 

2.  A   conceptacle  cut  in   meridional  direction  showing  the  tetra- 
sporangia.     Zeiss  2  x  BB. 

Figure  3.      Cheilosporum  californicum  (Dene.). 
A  portion  of  branch  showing  the  conceptacles.     The  scars  of  broken 
conceptacles  are  seen  as  deep  excavations,      x  ca.  3. 

Figures  4-8.      Cheilosporum  frondescens  (Post,  et  Rupr. ). 

4.  f.  typica;  a  portion  of  frond,      x  ca.  4. 

5.  "          a  fertile  branch.      x  ca.  5. 

6  and  6a.  f.  polymorpha;  portion  of  frond,      x  ca.  3]^. 

7.  "          a  fertile  branch.      x  ca.  4. 

8.  Cross  section  of  a  fertile  articulus  of  f.  typica.     The  dotted  line 
indicates  the  boundary  between  the  cortical  part  and  the  medullary. 

Figures  9—10.      Cheilosporum  planiusculum  (Kiitz.). 

9.  f.  normalis.      X  ca.  5. 

10.  f.  regularis.      X  ca.  5. 

Figures  11—14.    Cheilosporum  MacMillani. 

ii-i2.  Portions  of  the  fertile  branches  showing  pyriform  concep- 
tacles taking  places  of  the  pinnules  (n),  or  one  or  more  immersed 
in  the  pinnules  (12).  X  ca.  2. 

13.  Portions  of  a  fertile  branch,   showing  wart-like  conceptacles, 
one  of  them  are  found  inserted  at  the  apex  of  a  pinnule,      x  ca.  2. 

14.  Meridional   section   of    a    pyriform    conceptacle,   showing    an 
antheridium  :    the  granular  mass  in  the  cavity  is  an  aggregation  of 
spermatozoids.     Zeiss  2  x  BB. 


722  MINNESOTA    BOTANICAL    STUDIES. 

Figure  15.  Corallina  officinalis  L.  var  chilensis  (Kiitz.).  Merid- 
ional section  of  a  conceptacle,  showing  the  tetrasporangia.  Zeiss 
2  x  BB. 

Figures  16— 17.    Corallina  Vancouver iensis. 

O  I 

1 6.  f.  typica. 

17.  f.  densa. 

Figures  18-19.    Corallina  aculeata. 

1 8.  A  portion  of  frond.      X  ca.  4. 

19.  A  young  frond  found  in  a  brackish  pool   high  above  the  tidal 
mark. 


VOL.  II. 


MINNESOTA 


R.    UCHIYAMA    ET    K.    YENDO.       PHOTO. 


NICAL  STUDIES. 


PART  VI. 


HELIOTYPE    CO.,    BOSTON. 


VOL.  II. 


MlNNE 


H.    UCHIYAMA    ET    K.    VENDO.       PHOTO. 


AL  STUDIES. 


PART  VI. 


HELIOTYPE    CO.,    BOSTON. 


VOL.  II. 


R.    UCHIYAMA    ET    K.    YENDO.       PHOTO. 


PART  VI. 


VOL.  II. 


MlNNESC 


R.    UCHIYAMA    ET    K.    VENDO.       PHOTO. 


STUDIES. 


PART  VI. 


•IELIOTYPE    CO.,    BOSTON. 


VOL.  II. 


MINNESOTA 


R.    UCHIYAMA    ET    K.    YENDO.       PHOTO. 


:AL  STUDIES. 


PART  VI. 


HELIOTVPE    CO.,    BOSTON. 


VOL.  II. 


MlNNESO 


14 


K    YENDO.    DEL. 


L  STUDIES. 


PART  VI. 


1 


v  /  V ' 

m 


v\ 


:l 


7 


17 


-• 


15 


16 


18 


HELK5TYPE    CO.,    BOSTON. 


XLI.    OBSERVATIONS    ON    PTERYGOPHORA. 


CONWAY    MAcMlLLAN. 


Among  the  larger  and  less  thoroughly  studied  kelps  of  the 
Pacific  coast,  Pterygophora  californica  Ruprecht  has  seemed 
worthy  of  some  attention.  A  fine  series  of  specimens  was  se- 
cured during  the  summer  of  1901  at  the  Minnesota  Seaside  Sta- 
tion on  the  west  coast  of  Vancouver  Island,  and  while  all  points 
in  the  anatomy  and  life-history  cannot  be  elucidated  from  the 
material  in  hand,  it  has  seemed  best  to  present  an  account  of 
such  structural  and  developmental  facts  as  have  been  observed. 

The  name,  Pterygophora  californica,  appears  for  the  first 
time  in  botanical  literature  in  Ruprecht's  Algen-Stamme,*  in 
1848.  In  this  work  no  description  is  given  of  the  genus  and 
species,  Ruprecht  contenting  himself  with  the  statement  that 
the  new  genus  is  intermediate  between  Capca  (Ecklonia)  and 
Alaria — a  point  of  view  which  has  been  but  slightly  modified 
by  later  study — and  that  it  will  be  elsewhere  described.  In 
this  work,  however,  there  is  given  some  account  of  the  anatomy 
of  the  stem  and  the  characteristic  annular  structure  seen  in  the 
cross  section  is  mentioned  in  the  following  phrase:  "  Alle 
(zuweilen  bis  9)  Ringe  deutlich  und  gleich  stark  sind."  It  is  in 
this  paper,  also,  that  Ruprecht  announces  the  presence  of  muci- 
lage ducts  in  Pterygophora^  an  observation  which  has  since 
been  questioned  by  Areschoug. 

The  formal  description  of  Pterygophora  californica,  together 
with  a  plate  which  leaves  something  to  be  desired,  is  presented 
by  Ruprecht  in  his  "Pflanzen  aus  dem  nordlichen  Theile  des 
Stillen  Oceans,"  |  published  four  years  later.  The  specimens 
upon  which  the  description  of  Ruprecht  was  based  were  col- 

*  Ruprecht,  F.  J.  Bemerkungen  ueber  den  Bau  und  das  Wachsthi'm  einiger 
grossen  Algen-Stamme.  Mem.  Acad.  Imp.  Sci.  Nat.  Pctcrsb.  6 :  64,  70.  1848. 

f  Ruprecht,  F.  J.  Neue  oder  umvollstandig  bekannte  Pflanzen  aus  dem 
nordlichen  Theile  des  Stillen  Oceans.  Mem.  Acad.  Imp.  Sci.  Nat.  Petersb.  7  : 
17-19  (73-75)-  *•  5,  8.  1852. 

723 


724  MINNESOTA    BOTANICAL    STUDIES. 

lected  by  Wosnessenski  in  the  vicinity  of  "  Ross"  (that  is,  near 
San  Francisco)  on  the  coast  of  California  in  July,  1840.  The 
material  communicated  to  Ruprecht  comprised  some  middle- 
sized  plants  and  some  younger  specimens,  but  did  not,  appar- 
ently, enable  him  to  see  either  the  sporelings  and  unilaminate 
stage  or  the  mature,  full-grown  form.  Consequently  his  meas- 
urements need  modification  and  the  specific  description  should 
be  revised  to  include  much  larger  plants.  Anderson  is  the  only 
one  who  has  written  on  the  plant  in  whose  account  a  knowledge 
of  full-sized  plants  is  indicated.  In  his  Natural  History  of 
Santa  Cruz  county  *  it  is  stated  that  the  stem  is  from  one  to  six 
feet  in  length  and  that  the  leaves  are  from  one  to  five  feet 
long,  "  all  without  a  midrib  except  the  central  one  into  which 
the  flattened  stem  seems  to  be  lost,  giving  it  the  look  of  a  mid- 
rib." 

It  was  from  Anderson  that  Areschoug  received  his  specimens 
of  PterygopJwra  upon  which,  together  with  those  of  Ruprecht 
sent  him  from  the  collections  of  the  Academy  at  St.  Petersburg, 
he  based  his  brief  study  as  set  forth  in  Observationes  Phycolo- 
gicae.f  In  this  paper  Areschoug  gives  in  compact  form  the 
generic  and  specific  description  and  adds  some  observations  and 
corrections  to  the  earlier  account  of  the  Russian  botanist. 
Areschoug  observes  that  the  number  and  nature  of  the  growth- 
rings  in  the  trunk  seem  difficult  to  define.  He  is  skeptical  con- 
cerning the  lacunas  muciferse,  although  he  retains  in  his  revised 
description  of  the  genus  the  phrase  "  ad  peripheriam  internam 
annuli  intimi  collocata,  interdum  biseriata,"  crediting  the  ob- 
servation to  Ruprecht.  He  reexamined  the  younger  specimens 
upon  which  Ruprecht's  original  account  was  based  and  gives  a 
condensed  description  of  them.  The  youngest  plant  seen  ap- 
pears to  have  had  a  stipe  20  cm.  long  and  2-5  mm.  broad  with 
a  lamina  35  cm.  long  and  6-7  cm.  broad.  As  will  be  seen  later 
my  own  plant,  "  B  ",  indicates  that  these  measurements,  based 
upon  dried  material,  need  correction.  Areschoug  received  mate- 
rial from  "  Vera  Cruz,"  California  (meaning  Santa  Cruz),  sent 
him  by  Dr.  Anderson  and  notes,  doubtless  upon  the  authority  of 
Anderson,  that  the  plant  grows  also  at  San  Francisco.  In 

*  Anderson,  C.  I/.  The  natural  history  of  Santa  Cruz  county,  Oakland,  Cal., 
24.  No  date.  (1892?) 

t  Areschoug,  J.  E  Observationes  Phycologicae,  Part  5:  De  Latninariaceis 
Nonnullis,  n.  1884. 


MacMillan:  OBSERVATIONS  ON  PTERYGOPHOKA.  725 

Anderson's  List  of  California  marine  algae,*  Pterygophora  cali- 
f arnica  is  recorded  as  "  common  or  occasional  to  all  the  Cali- 
fornia coasts."  The  southernmost  record  of  its  occurrence  that 
I  have  happened  to  find  is  in  Orcutt's  Flora  of  Southern  and 
Lower  California,  f  The  plant  is  here  credited  to  San  Diego, 
based  upon  collections  by  Daniel  Cleveland.  The  northernmost 
point  from  which  specimens  have  been  taken  seems  to  be  Port 
Renfrew,  Vancouver  Island,  B.  C.,  giving  the  plant  a  range  of 
over  16°  of  latitude.  It  very  probably,  however,  extends  up 
the  Alaskan  coast.  In  some  observations  upon  the  distribution 
of  Pterygophora,  Setchell  J  notes  that  the  characteristic  Lami- 
narieas,  Agarieae  and  Alarieae  of  the  North  Pacific  "  stop  at 
about  Puget  Sound  which  is  the  terminus  of  the  isothere  of  15°, 
but  Costaria  turnert'Grev.  and  Alaria  esculenta  (L.)  Grev.  con- 
tinue to  Monterey,  nearly  to  the  20°  line,  although  they  are 
found  only  at  '  exposed  points.'  .  .  .  An  interesting  case  is  that 
of  Pterygophora  californica  Rupr.  which  is  reported  by  Dr.  C. 
L.  Anderson  as  growing  at  Monterey  all  the  year  round,  but  is 
reported  by  Mr.  Daniel  Cleveland  as  occurring  at  San  Diego 
only  from  February  until  May  and  in  deep  water."  This  state- 
ment seems  to  be  based  upon  the  account  of  Pterygophora  given 
by  Hervey§  in  his  Sea  Mosses  in  the  preparation  of  which  he 
had  the  assistance  of  Dr.  Anderson.  Pterygophora,  therefore, 
seems  to  have  a  somewhat  wider  range  than  the  majority  of  the 
North  Pacific  Laminariacese.  The  plants  collected  at  Port  Ren- 
frew were  so  abundant  and  robust  that  I  am  inclined  to  think 
that  the  region  of  maximum  development  may  be  along  the 
British  Columbian  rather  than  along  the  Californian  coast. 
Ruprecht's  plants  were  Californian  ;  those  of  Areschoug  were 
from  the  vicinity  of  Monterey,  as  was  also  the  specimen  of 
Hervey  :  the  specimens  of  Cleveland  from  San  Diego  do  not 
seem  to  have  been  recorded  as  of  unusual  size.  The  plants  of 
Port  Renfrew,  some  of  them  with  trunks  nearly  three  inches  in 
diameter  and  eight  feet  in  length,  exceed  the  recorded  measure- 
ments and  indicate  thus  a  particularly  luxuriant  growth  in  that 

*  Anderson,  C.  L.  List  of  California  marine  algae,  with  notes.  Zoe,  a  :  220. 
1891. 

t  Orcutt,  C.  R.     Flora  of  Southern  and  Lower  California,  13.     1885. 

|  Setchell,  W.  A.  On  the  classification  and  geographical  distribulion  of  the 
Laminariaceae.  Trans.  Conn.  Acad.  g  :  370.  1893. 

\  Hervey,  A.  B.  Sea  Mosses.  A  collector's  Guide  and  an  introduction  to 
the  study  of  marine  algae,  88.  1881. 


726  MINNESOTA    BOTANICAL    STUDIES. 

region.  The  measurements  given  by  different  authors  are  as 
follows  :  Ruprecht  (1.  c.),  stipe  6-9  inches  long,  laminae  up  to  2 
feet  long :  Areschoug  (1.  c.),  stipe  30  cm.  long,  lamina  up  to 
one  meter  in  length  ;  De  Toni,*  stipe  30  cm.  long,  lamina  up 
to  one  meter  in  length  (measurements  evidently  quoted  from 
Areschoug) ;  Hervey  (1.  c.),  stipe  2  or  3  feet  long,  lamina  2  feet 
or  more  long ;  Setchell  (1.  c.),  stipe  i  to  2  feet  long,  measure- 
ment of  leaves  not  given;  Anderson  (1.  c.),  stipe  i  to  6  feet 
long,  lamina  i  to  4  feet  long.  Of  these  measurements,  Ander- 
son's is  the  only  one  that  is  approximately  correct  for  the  aver- 
age plant  as  observed  on  the  Straits  of  Fuca. 

The  first  specimen  of  Pterygophora  seen  on  the  Vancouver 
coast  was  a  battered  and  eroded  stem  which  had  been  cast  up 
by  the  tide.  It  was  between  six  and  seven  feet  in  length  and 
2.5  inches  in  thickness  near  the  base.  Later  another  specimen, 
not  quite  so  large,  was  extracted  from  a  pile  of  wrack  at  the  head 
of  a  little  cove  and  this  had  a  few  dilapidated  leaves  still 
attached.  Examination  of  the  shore  yielded  several  speci- 
mens, some  of  which  were  in  an  excellent  state  of  preserva- 
tion, but  a  few  days  later  some  growing  beds  were  discovered 
and  the  plant  was  observed  in  more  detail.  Its  selection  of  an 
habitat  is  interesting.  A  favorite  place  for  its  development 
seemed  to  be  on  the  bottom  of  deep,  narrow  chasms  in  which 
there  was  from  twelve  to  fifteen  feet  of  water  at  low  tide.  It 
occurred  abundantly  on  the  bottom  of  a  circular  hole  commu- 
nicating with  the  sea  by  a  narrow  deep  inlet  and  exposed  to 
heavy  surge.  It  was  afterwards  found  that  this  was  its 
characteristic  position  and  that  it  habitually  came  closer  to  the 
rocks  than  either  Nereocystis  or  Macrocystis.  It  preferred 
stations  where  the  water  was  constantly  in  motion  and  did  not 
seem  so  abundant  in  quiet  coves.  As  a  surge  plant  it  grew 
lower  than  Lessonia  and  it  may,  perhaps,  be  described  as  oc- 
cupying the  lowest  position  of  the  surge  kelps  along  this  coast. 
To  this  precise  locality  the  plant  shows  certain  structural  adapta- 
tions. The  holdfast  is  massive,  enabling  it  to  cling  firmly  to  the 
rocks,  notwithstanding  the  strong  movement  of  the  sea.  The 
stem  is  exceedingly  stout — being  indeed  one  of  the  strongest 
algal  structures  known — and  is  capable  of  resisting  great 
tensile  strain.  While  not  particularly  elastic  it  is  bent  from 
side  to  side  without  difficulty  or  damage  to  its  structure.  The 

*  De  Toni,  J.  B.     Sylloge  Algarum,  3  :  352.     1895. 


MacMillan:  OBSERVATIONS  ON  PTEKYGOPHORA.  727 

long  leaves,  often  as  many  as  forty  in  the  tuft,  hang  down 
beside  the  stem  and  as  the  plant  bends  from  side  to  side  they 
are  swept  along  the  bottom,  thus  accounting  for  the  erosion 
of  their  ends  so  characteristic  of  this  species.  The  central 
lamina  is  invariably  eroded,  and  only  the  younger  and  shorter 
pinnae  are  perfect,  all  the  older  ones  having  lost  their  tips 
through  the  constant  brushing  back  and  forth  on  the  rocky 
bottom.  The  relative  lengths  of  the  leaves  and  of  the  stem  are 
regulated  by  this  habit  of  the  growing  plant  and  where  the 
surge  was  most  violent  plants  were  to  be  found  with  compara- 
tively long  stems  and  short  leaves,  but  where  the  surge  was 
less  violent  the  leaves  and  stem  were  more  nearly  the  same 
length,  or  the  leaves  might  some  of  them  even  exceed  the 
stem. 

The  general  appearance  of  young  Pterygophora  plants  as 
seen  upon  the  bottom  is  not  unlike  that  of  Nereocystis.  Their 
attitudes  with  the  erect  stem  and  the  dependent  leaves  are  very 
similar.  The  older  Pttrygopkora  plants,  from  their  much 
more  massive  stem  and  shorter  leaves,  can  be  distinguished  at 
a  glance. 

In  order  to  collect  an  abundant  series  of  Pterygophora  calif or- 
nica  use  was  made  of  a  tool  which  may  be  described  as  a  combi- 
nation of  chisel  and  hook  on  the  end  of  a  long  slender  pole,  by 
which  the  holdfasts  were  cut  and  the  plant  dragged  to  the  sur- 
face. In  this  way  a  sufficient  quantity  of  material  was  ob- 
tained from  which  four  plants  of  different  ages  are  selected  for 
description. 

Plant  "  A."  This  is  the  youngest  specimen  seen.  It  meas- 
ures 12  mm.  in  length,  of  which  the  stipe  and  primitive  disk 
constitute  but  2  mm.,  the  rest  being  lamina.  In  this  plant  the 
lamina  is  already  eroded  distally.  It  measures  5  mm.  across  at 
its  broadest  part  and  narrows  down  abruptly  to  the  stipe,  which 
is  5  mm.  in  diameter.  The  primitive  disk,  almost  exactly  cir- 
cular in  shape,  measures  2  mm.  in  diameter.  At  first  the  growth 
of  the  stipe  in  length  is  decidedly  slow,  but  when  the  lamina 
has  become  about  20  mm.  in  breadth  the  stipe  begins  to  elon- 
gate. In  plants  under  30  mm.  in  length  the  poorly  defined 
midrib  of  Pterygophora  has  not  begun  to  develop  and  the  lam- 
ina seems  perfectly  homogeneous  throughout.  In  this  respect 
the  plant  is  in  marked  contrast  with  Alaria  sporelings,  for  in 
them  the  midrib  will  have  already  strongly  developed  in  plants 


728  MINNESOTA    BOTANICAL    STUDIES. 

of  corresponding  size.  The  young  forms  of  Pterygophora 
californica  show  the  characteristic  shape  and  developmental 
sequence  of  the  Laminariaceae,  resembling  particularly  young 
plants  of  Laminaria  saccharina. 

Plant  "  B."  This  specimen  was  collected  by  Mr.  K.  Yendo 
and  was  kindly  presented  to  me  by  him.  It  is  apparently  of  about 
the  same  age  as  Ruprecht's  youngest  plant.  The  whole  plant 
is  35  cm.  in  length,  of  which  10  cm.  is  holdfast  and  stipe, 
and  25  cm.  lamina.  The  lamina  is  8  cm.  in  breadth  at  its 
broadest  part.  The  tip  is  eroded,  as  usual,  and  in  this  particu- 
lar specimen  the  margin  is  imperfect.  The  stipe  is  8  mm.  in 
diameter,  3  cm.  below  the  base  of  the  lamina,  and  5  mm.  in  di- 
ameter just  above  the  holdfast,  where  it  is  circular  in  cross-sec- 
tion. It  is,  however,  elliptical  in  cross  section  near  the  base  of 
the  lamina,  the  stipe  being  flattened  in  the  plane  of  the  lamina. 
In  this  specimen  the  midrib  is  beginning  to  differentiate  and  is 
well-marked  for  a  distance  of  5  cm.  above  the  base  of  the  lam- 
ina and  faintly  marked  for  10  cm  farther  towards  the  tip. 
Distally  it  quite  disappears.  Certain  hapteric  outgrowths  are 
decidedly  long  and  slender  in  this  plant — much  more  developed 
than  ordinarily. 

In  plant  "  B,"  less  than  i  cm.  below  the  base  of  the  lamina, 
are  seen  two  small  emergences,  opposite  each  other  on  the  sides 
of  the  stipe.  These  are  the  growing  points  destined  to  produce 
the  first  pair  of  pinnae. 

Plant  "  C."  This  is  a  somewhat  older  individual  in  which 
twelve  pinnae  have  been  developed.  The  whole  plant  is  45  cm. 
in  length  from  the  holdfast  to  the  eroded  tip  of  the  central 
lamina.  From  the  holdfast  to  the  lowest  pinna  is  15  cm.  In 
this  plant  the  four  lower  pinnae  are  of  a  deep  chocolate  brown 
color  and  very  much  eroded  and  perforated.  They  present 
some  points  of  anatomical  interest  as  will  be  indicated  later. 
The  upper  pinnae  are  olive  brown  in  color  and  the  four  upper- 
most have  perfect  tips,  characteristically  rounded,  giving  to  the 
whole  young  pinna  a  distinctly  spatulate  shape.  In  texture 
these  young  pinnae  in  the  fresh  plant  are  quite  unlike  the  sporo- 
phylls  of  Alaria  with  which  they  have  been  compared.  To  the 
touch  they  feel  not  unlike  thin  sheets  of  celluloid.  The  central 
lamina  has  a  more  leathery  feel,  like  Laminaria  or  Lessonia. 

In  this  plant  the  stipe  is  8  mm.  in  diameter,  midway  between 
the  holdfast  and  the  pinnae.  In  the  region  of  pinnae  it  is  de- 


MacMillan:  OBSERVATIONS  ON  PTERYGOPHORA.  729 

cidedly  flattened,  measuring  9  mm.  in  width  by  3  mm.  in  thick- 
ness. The  pinnae  do  not  stand  always  directly  opposite  each 
other,  although  this  in  general  is  their  position.  Abortion  of 
one  pinna  of  the  pair  may  be  observed  in  this  plant  at  two  points. 
The  base  of  the  pinnae  is  more  attenuate  than  that  of  the  cen- 
tral lamina.  The  holdfast  is  partly  cut  away,  but  is  flattened 
out  and  compacted  much  more  than  in  plant  B. 

Plant  "  D."  This  plant  was  collected  in  July,  preserved  in 
formalose,  and  brought  to  Minneapolis  for  study.  It  was  found 
growing  with  several  others  about  twelve  feet  below  the  surface 
of  the  water  at  low  tide.  The  stipe  from  the  holdfast  to  the 
lowest  pinnae  is  2  meters  in  length.  From  the  lower  pinnae  to 
the  base  of  the  central  lamina  is  i  dm.  while  the  central  lamina 
is  1.5  meters  in  length.  On  each  side  of  the  stipe,  extending 
along  its  margin  for  a  little  less  than  a  decimeter,  are  the  tufts 
of  lateral  pinnae,  twenty  on  each  side.  The  longest  pinna  with 
uneroded  end  measures  i  meter,  but  pinnae  with  eroded  ends 
are  present,  1.5  meters  in  length.  The  breadth  of  the  central 
lamina  is  i  dm.,  the  midrib  being  4  cm.  broad.  The  broadest 
pinna  measures  7.5  cm.  from  margin  to  margin.  All  margins 
of  full  grown  pinnae  are  undulate.  This  character  is  especially 
marked  in  the  central  lamina.  One  difference  between  a  plant 
of  the  age  of  "  D"  and  a  younger  form  such  as  "  C"  lies  in 
the  distance  between  the  adjacent  pairs  of  pinnae.  In  plant 
"  C,"  for  example,  the  upper  pinnae  are  three  cm.  apart  along 
the  stipe  and  this  character  is  also  indicated  in  Ruprecht's 
plate.  In  an  older  plant,  such  as  "D, "the  pinnae  are  very 
much  crowded  together,  so  much  so,  indeed,  that  they  crowd 
each  other  out  of  a  strictly  marginal  position.  The  fully  de- 
veloped pinnae,  in  "  D  "  are  all  massed  within  a  linear  distance 
of  5  cm.,  while  in  "  C  "  they  are  distributed  over  twice  as  much 
space. 

The  stipe  in  this  specimen  is  5  cm.  in  diameter,  2  dm.  above 
the  holdfast.  Near  the  holdfast  it  is  i  dm.  in  diameter.  Nearer 
the  pinnae  it  becomes  flattened  in  cross  section,  first  appearing 
as  elliptical,  then  as  lenticular,  the  edges  becoming  sharp  2  dm. 
below  the  lowest  pinnae.  Along  the  sharp  edges  the  scars  ol 
pinnae  which  have  been  sloughed  off  are  abundant.  The  stipe 
in  the  region  of  pinna  attachment  is  3  cm.  broad  and  7.5  mm. 
thick.  The  stipes  of  the  full-grown  pinnae  are  4  mm.  in  diameter 
and  the  base  of  the  central  lamina  is  i  cm.  broad  and  3  mm.  in 


730  MINNESOTA    BOTANICAL    STUDIES. 

thickness.  A  transsection  through  the  stipe  of  this  plant  shows 
24  concentric  rings  of  growth  not  all  of  which  were  of  equal 
thickness.  The  pith  at  the  center  has  the  lenticular  outline 
characteristic  of  arborescent  Laminariaceae  and  was  8  mm.  by 
2  mm.  in  cross  section.  The  pith  occupies  a  greater  portion  of 
the  cross  section  nearer  the  region  of  pinnae.  Just  below  the 
pinna  scars  a  cross  section  showed  it  to  be  3  cm.  in  length  by  3 
mm.  in  width. 

From  the  measurements  of  this  plant,  by  no  means  the  largest 
seen,  it  becomes  apparent  that  the  size  of  Pterygophora  cali- 
fornica  has  been  much  underestimated,  previous  descriptions  of 
it  having  been  made  from  immature  material.  There  seems  to 
be  no  reason  to  doubt  that  this  plant  is  perennial.  It  gives  every 
structural  indication  of  persisting  for  a  series  of  years  and  re- 
plenishing its  pinnae  with  the  recurring  seasons.  Its  massive 
stipe  and  the  base  of  the  central  lamina  survive  the  winter 
storms  and  in  the  spring  fresh  pinnae  are  produced  and  the 
central  lamina  is  extended  by  the  well-known  basal  growth 
characteristic  of  the  family  to  which  it  belongs.  In  this  way, 
doubtless,  very  large  plants  may  develop.  There  is  one  frag- 
ment in  our  collection  comprising  the  pinna  region  of  a  plant 
which  by  comparative  measurements  must  have  been  four  meters 
or  more  in  length.  The  flattened  stipe  between  the  pinnae  is  6 
cm.  broad  and  the  base  of  the  central  lamina  measures  3  cm. 
in  width. 

The  relation  of  the  genus  Pterygophora,  to  the  other  genera 
of  the  Laminariacese  has  been  a  matter  of  some  uncertainty. 
Ruprecht  regarded  it  as  intermediate  between  Ecklonia  and 
Alaria.  Agardh*  associates  Pterygopkora  with  Alaria.  Ares- 
choug  takes  it  up  between  Lessonia  and  Ecklonia,  but  this  per- 
haps can  scarcely  be  regarded  as  an  expression  of  his  opinion 
regarding  its  true  position.  By  the  older  systematists  the  genus 
has  been  connected  closely  with  Laminaria  and  it  occupies  a 
position  next  to  Laminaria  in  De  Toni's  Sylloge  Algarum  and 
also  in  Kjellman's  Laminariaceae  (1.  c.)  where  Pterygophora  is 
placed  between  Laminaria  and  Ecklonia  in  Tribe  VI.,  Lami- 
narieae.  Setchell  (1.  c.)  connects  the  genus  with  Alaria  under 
the  Tribe  Alariideae.  The  mid-lamina  of  Pterygophora  is 
strongly  suggestive  of  certain  species  of  the  genus  Laminaria, 
so  much  so  that  when  Areschoug  described  Laminaria 

*  Agardh,  J.  G.     De  Laminarieis,  Lund  Univ.  Arsskr.  4:  i. 


Ma cM Ulan:  OBSERVATIONS  ON   PTERYGOPHORA.  731 

japonica,*  Agarth  f  suggested  that  the  type  specimen  was 
nothing  other  than  a  mid-lamina  of  Ptcrygophora.  While  this 
notion  of  Agardh's  was  incorrect,  a  comparison  of  specimens  or 
reference  to  Suringar'sJ  plate  of  Laminaria  japonica  will  make 
it  clear  how  natural  might  have  been  such  a  supposition  on  his 
part.  One  also  finds  in  the  genus  Laminaria  forms  suggesting 
the  pinnate  disposition  of  laminae  in  Ptcrygophora.  Such  a 
plant  is  figured  and  described  by  Kjellman§  under  the  name  of 
Laminaria  radicosa.  In  this  plant  lateral  outgrowths  occur 
upon  the  stipe  below  the  lamina  in  quite  the  same  position  in 
which  they  are  developed  in  Ptcrygophora.  They  are  not, 
however,  functional  as  additional  laminae,  nor  do  they  particu- 
larly increase  the  photosynthetic  vigor  of  the  plant.  Laminaria 
radicosa  may,  nevertheless,  be  regarded  perhaps  as  showing  a 
transition  to  the  type  of  Ptcrygophora. 

There  are  some  objections  to  the  classification  of  Plcrygo- 
phora  with  Alaria.  Among  these,  the  character  of  the  young 
plant  should  be  given  weight.  In  Alaria  the  midrib  is  dif- 
ferentiated at  an  early  stage  and  is  exceedingly  distinct  in  plants 
only  two  centimeters  in  length,  while  in  Pterygophora  plants 
35  centimeters  in  length  show  the  midrib  but  indistinctly  in 
the  basal  portion  of  the  lamina.  Anatomically  Pterygophora 
conforms  to  the  type  of  the  Laminarieae  in  the  general  character 
of  its  tissues,  differing  in  some  marked  particulars  from  Alaria, 
although  resembling  the  latter  in  absence  of  mucilage  canals — 
structures  which  are  present  in  most  species  of  Laminaria. 
The  distinction  of  outer  and  inner  cortex  which  is  not  always  to 
be  made  out  in  Alaria  is  very  clear  in  Pterygophora.  Upon 
the  whole  there  would  seem  to  be  little  objection  to  the  classifi- 
cation of  Plcrygophora  in  the  tribe  Laminarieae.  Taking  every- 
thing into  account,  however,  it  will  perhaps  be  best  to  consider 
the  genus  as  transitional  between  the  Laminarieae  and  the 
Alariideae. 

An  examination  of  the  anatomy  of  Pterygophora  seems 
further  to  strengthen  the  view  of  its  close  relation  to  Laminaria, 
while  the  differentiation  of  its  organs  no  doubt  makes  it  readily 


*  Areschoug,  J.  E.     Phyceae  Capenses,  29. 

t  Agardh,  J.  G.     Proc.  Soc.  Phys.  Lund.  Bot.  Notiser,  1883  :  108.     1883. 
JSuringar,  W.  F.  R.     Algae  Japonicae,  pi.  //.     1870. 

g  Kjellman,  F.  R.,  and  Peterson,  J.  V.     Om  Japans  Laminariaceer.    Ur  Vcga- 
Expeditionens  Vetensk.  lakttagelser. ,  4:  259,  pi.  10.     1885. 


732  MINNESOTA    BOTANICAL    STUDIES. 

comparable  with  Alaria.  The  structure  of  the  stipe  differs 
decidedly  from  that  of  Lessonia  which  I  have  previously  ex- 
amined.* In  both  genera  there  are  strongly  marked  growth- 
rings  which,  as  will  be  seen,  do  not  arise  in  precisely  the  same 
way.  A  detailed  account  of  the  anatomy  follows.  From  it 
some  notion  may  be  derived  of  the  histological  interrelation  of 
Ptcrygophora,  Laminar ia  and  Lessonia. 

The  most  important  literature  on  the  anatomy  of  the  Lamina- 
riaceae  has  been  previously  cited  f  and  it  will  not  be  necessary 
to  refer  to  it  further  at  this  time  except  as  some  particular  point 
may  require  elucidation.  To  the  papers  of  Wille,  Grabendorfer, 
Reinke,  Rosenthal,  Oliver,  Ruprecht  and  others  students  are 
indebted  for  researches  which  have  laid  the  foundation  for  a 
knowledge  of  the  anatomy  of  the  Laminariaceae. 

The  holdfast. — The  study  of  this  structure  as  of  the  other  or- 
gans of  Pterygophora  is  based  upon  a  series  of  slides  prepared 
from  material  collected  at  the  Minnesota  Seaside  Station, 
killed  in  chromic  acid,  and  transferred  into  70  per  cent,  alcohol 
in  which  condition  it  was  brought  to  Minneapolis  for  study. 
Most  of  the  sections  have  been  cut  freehand,  treated  with  various 
reagents  and  stains  and  mounted  in  glycerine-jelly.  Russow's 
callus  reagent,  chlor-zinc  iodide  and  a  variety  of  stains,  includ- 
ing particularly  the  fuchsin  and  iodine-green  combination  and 
aniline  water  safranin,  have  been  employed  to  bring  out  de- 
tails of  structure. 

The  primitive  disc  shows  no  points  of  special  interest,  not 
differing  particularly  in  structure  from  that  already  described 
for  Ncreocystis,\  nor  at  first  do  the  hapteric  branches  in  their 
origin  and  structure  show  characters  worthy  of  especial  com- 
ment. The  haptere  originates  through  the  activity  of  a  circu- 
lar cambial  area  at  the  edge  of  the  primitive  disc  or  from  the 
lower  rhizogenous  area  of  the  stipe.  Callosities  on  the  stipe, 
such  as  those  described  for  Nercocystis  and  there  believed  to  be 
equivalent  to  hapteric  branches,  have  not  been  discovered  in 
Ptcrygopkora,  though  on  one  specimen  some  curious  gall-like 
swellings,  doubtless  teratological  or  pathological  in  their  nature, 
were  observed.  The  numerous  hapteric  outgrowths  of  Ptcry- 

*  MacMillan,  C.  Observations  on  Lessonia,  Bot.  Gaz.  30:  318.  pi.  19-21. 
1900. 

t  MacMillan,  C.     1.  c. 

%  MacMillan,  C.  Observations  on  Nereocystis,  Bull.  Torr.  Club,  26:  273. 
pi.  j6f,  362.  1899. 


MacMittan:  OBSERVATIONS  ON  PTERYGOPHORA.  733 

gophora  branch  dichotomously  arid  build  a  strong  holdfast  re- 
sembling that  of  Lessonia  rather  than  that  of  Nercocystis. 
Each  young  hapteric  branch,  in  cross  section,  shows  the  char- 
acteristic structure,  a  great  central  mass  of  parenchymatous 
tissue  surrounded  by  an  ill-defined  cortical  area  with  an  hypo- 
dermal  cambium.  No  pith  is  present  and  the  growth  in  length 
and  thickness  of  the  hapteres  proceeds  solely  by  cambial  ac- 
tivity. In  older  holdfasts  distinct  growth-rings  appear — some- 
thing that  was  not  seen  in  the  holdfasts  of  Lessonia,  and  seems 
not  to  occur  in  the  hapteric  branches  of  Nereocystis,  the  indi- 
viduals of  which  are  shorter-lived.  The  appearance  of  these 
rings  of  growth  in  the  secondary  cortical  tissues  of  the  holdfast 
seems  to  be  due  rather  to  rhythmic  changes  in  the  character  of 
the  cell  contents  than  to  regular  successions  of  larger  and 
smaller  cells,  concentrically  arranged.  As  will  appear,  this 
character  serves  to  distinguish  to  some  extent  between  the 
growth-rings  of  the  holdfast  and  of  the  stipe.  In  both  organs 
the  elements  of  the  secondary  cortex  are  arranged  in  extremely 
regular  rows,  as  seen  in  cross  section.  This  regularity  of  ar- 
rangement does  not  extend  to  the  primary  parenchymatous 
tissue  of  the  hapteric  branch,  so  that  the  appearance  of  a  cross 
section  of  the  older  hapteres  may  be  described  as  follows :  At 
the  center  is  a  large  more  or  less  circular  group  of  parenchy- 
matous elements  regularly  hexagonal  in  outline,  varying  in  size 
between  rather  narrow  limits.  Towards  the  periphery  this  cen- 
tral tissue  imperceptibly  merges  with  the  secondary  tissue,  the 
cells  of  which  become  more  quadrate  in  outline  and  assume  the 
characteristic  position  in  rows  which  can  be  followed  without 
break  directly  to  the  cambial  zone  which  lies  near  the  periphery 
of  the  organ.  In  the  secondary  tissue  there  are  numerous  rings 
of  growth  and  the  cross  section  of  an  old  haptere,  a  centimeter 
or  more  in  diameter,  looks  not  unlike  a  section  of  stipe,  save 
for  the  absence  of  the  characteristic  lenticular  pith.  This  is 
altogether  wanting  in  the  hapteres.  The  growth-rings  do  not, 
however,  appear  to  arise  consistently  through  quite  the  same 
anatomical  conditions  as  those  of  the  stipe.  Well-marked  rings 
in  the  holdfast  may  exist  without  difference  in  the  size  of  the 
elements  of  which  they  are  composed.  The  optical  appear- 
ance, therefore,  is  in  all  hapteres  examined  determined  by  dif- 
ference in  the  cell  contents.  A  zone  of  cells  will  be  formed  in 
which  the  contents  seem  to  be  more  dense.  Outside  a  zone,  in 


731  MINNESOTA    BOTANICAL    STUDIES. 

which  the  contents  are  less  dense,  will  appear,  and  this  alter- 
nation continues  even  into  the  cambial  zone  where  single  layers 
of  cells  will  be  found  with  the  denser  contents  alternating  with 
layers  showing  the  opposite  appearance.  Besides  this  differ- 
ence others  have  not  been  discovered  to  account  for  the  ringed 
appearance  of  old  hapteres. 

As  to  the  substance  in  the  cells  which  by  its  greater  density 
gives  the  ringed  appearance  it  does  not  seem  to  differ  from  the 
material  which  has  been  studied  by  a  number  of  observers, 
especially  in  the  Fucaceas.  It  has  been  described  by  Reinke* 
as  a  fatty  oil,  in  which  view  Hansen  f  practically  coincides  and 
considers  that  the  Phaeophycea?  in  general  produce  fat  instead 
of  starch  by  their  assimilation.  The  same  material,  however, 
has  been  described  as  Phaeophyceenstarke  by  Schimper,  as 
fucosan  by  Schmitz  and  Hansteen,  as  showing  a  tannin  reac- 
tion by  Berthold,  as  phloroglucin-containing  material  by  Bruns, 
as  connected  with  physodes  by  Crato  and  as  polysaccharids,  in 
constitution  allied  to  mucine,  by  Koch.  In  the  holdfast  of 
Pterygophora  the  material  sometimes  fills  the  whole  cell  with  a 
homogeneous  refringent  mass,  which  in  the  denser  parts  of  the 
ring  has  decidedly  the  same  appearance  optically  that  is  shown 
by  the  polysaccharid  granules  of  the  lamina  and  stipe  to  be  de- 
scribed later.  In  other  instances  the  refringent  bodies  may  be 
distinguished  from  a  generally  granular  protoplasmic  slime 
which  encloses  them.  Without  going  into  the  disputed  ques- 
tion of  the  true  chemical  character  of  the  cell-contents  of  Lami- 
nariaceae,  it  may  be  said  that  the  ringed  appearance  of  hapteric 
branches  in  Pterygophora  is  due  to  the  alternately  more  vigorous 
and  less  vigorous  production  of  certain  substances  connected 
with  the  assimilative  processes  of  the  plant.  These  substances 
occur  in  the  stipe  and  lamina  as  well  as  in  the  hapteres,  but  are 
there  not  invariably  the  cause  of  a  ringed  appearance,  being 
disposed  in  special  cells  without  any  apparent  reference  to  the 
rhythm  of  secondary  growth. 

The  stipe. — Sections  were  taken  first  from  plant  "  B,"  the  uni- 
laminate  stage  and  then  from  mature  plants.  In  none  of  them 
could  the  mucilage  ducts  of  Ruprecht  be  discovered  and 


*  Reinke,  J.  Beitraege  zur  Kenntniss  der  Tange.  Pringsh.  Jahrb.  fur  wis- 
sensch.  Bot.  10  :  317.  1876. 

t  Hansen.  Ueber  Stoffbildung  bei  den  Meeresalgen.  Milth.  Zool.  Stat. 
Neap.  IT  :  276. 


MacMillan :  OBSERVATIONS  ON  PTERYGOPHORA  735 

Pterygo-phora  may  safely  be  described  as  devoid  of  these 
canals.  The  cross  section  of  a  young  stipe  shows  the  character- 
istic lenticular  pith-web,  composed  of  anastomosing  filaments 
with  numerous  trumpet  hyphae  intermingled.  Chloroplasts 
are  abundant  in  this  tissue  and  occur  more  or  less  sparingly  in 
the  perimedulla.  Surrounding  the  pith  one  finds  the  cells  of 
the  cortical  tissue  very  regularly  hexagonal  in  shape,  arranged 
in  remarkably  perfect  radial  rows  and  diminishing  gradually  in 
size  towards  the  periphery.  Chloroplasts  are  absent  from  most 
of  the  cells  of  this  tissue,  but  appear  again,  in  the  smaller 
cells  near  the  periphery.  At  about  the  depth  at  which  chloro- 
phyll becomes  abundant  the  tissue  is  lacuniferous  and  the  outer 
cortex  readily  separates  from  the  inner.  The  cells  of  the 
outer  cortex  are  generally  not  hexagonal,  but  cambial  condi- 
tions cause  them  to  assume  the  rectangular  outline  in  cross 
section.  The  small  densely  colored  cells  of  the  epidermis  and 
hypodermis  are  uniformly  quadrangular.  Longitudinal  sec- 
tions through  material  of  this  age  show  the  inner  cortex  to  be 
made  up  of  prosenchymatous  elements  not  pitted  or  armed  and 
the  walls  comparatively  thin  in  the  region  near  the  pith,  but 
becoming  thicker-walled  and  beginning  to  present  the  pitted 
structure  closer  to  the  periphery.  The  cells  of  the  outer  cortex 
seem  to  have  a  special  capacity  for  dividing  transversely  and 
periclinally  in  young  material,  but  in  older  stipes  they  divide 
radially  with  equal  ease.  In  mature  stipes  the  extraordinarily 
regular  radial  rows  of  cells  seen  in  cross  sections  may  be  ob- 
served to  originate  from  rows  of  cambial  cells  which  have 
divided  radially  in  the  outer  cortex  and  have  there  established 
the  general  radial  arrangement  of  the  tissues. 

Sections  through  the  mature  stipe  show  a  structure  of  the 
organ  in  cross  section  reminding  one  very  much  of  the  tracheids 
and  their  arrangement  in  the  Coniferae.  The  pits,  however, 
are  not  upon  the  radial  faces  of  the  elements,  but  upon  the  con- 
centric. The  cells  are  all  of  about  the  same  size  and  stand  in 
rows  radiating  in  a  most  regular  fashion  from  the  pith  to  the 
circumference.  There  is  often  not  the  slightest  difficulty  in 
observing  that  the  appearance  of  growth-rings  is  due  to  the 
gradual  diminution  in  the  diameter  of  the  cells  until  they  have 
become  distinctly  flattened,  followed  abruptly  by  the  production 
of  cells  of  slightly  larger  lumina.  That  is  to  say,  the  occasion 
for  the  ringed  appearance  of  the  stipe  is  structurally  quite  com- 


736  MINNESOTA    BOTANICAL    STUDIES. 

parable  with  that  condition  so  fully  studied  in  the  stems  of 
Coniferas  and  woody  Dicotyledons.  In  other  instances,  how- 
ever, the  rings  in  the  stipe  seem  to  arise  quite  as  in  the 
hapteres. 

There  is  in  the  stipe  of  Pterygophora  a  plain  distinction  be- 
tween the  first  or  primary  structure  of  the  cortex  and  the  second- 
ary structure  which  is  established  in  the  lacuniferous  period  of 
growth.  It  is  at  this  time,  during  the  first  year,  that  the  outer 
cortex  readily  peels  from  the  inner.  Later,  with  the  resump- 
tion of  growth  in  thickness,  the  lacunae  are  filled  by  the  radially 
dividing  cambial  tissue.  The  tissue  of  the  first  or  innermost 
ring,  surrounding  the  pith,  differs  in  appearance  from  that  of 
subsequent  rings.  The  cells  diminish  in  diameter  towards  the 
periphery  of  the  ring  until  they  have  the  look  of  stereome  in 
cross  section.  In  this  part  of  the  ring  the  cells  are  very  strongly 
pitted,  in  marked  contrast  to  the  inner  cells  of  the  primary  cor- 
tex where  the  pits  are  but  occasional.  All  the  cells,  however, 
of  the  secondary  cortex  as  displayed  in  subsequent  growth- 
rings  are  strongly  pitted.  Another  distinction  between  the  cells 
of  the  innermost  ring  and  those  of  subsequent  rings  is  that  the 
elements  of  the  first  ring  are  slenderer,  more  prosenchymatous, 
almost  approaching  the  fibrous  shape  peripherally,  while  the 
elements  of  later  rings  are  shorter,  more  parenchymatous  and 
not  at  all  suggestive  of  fibrous  tissue. 

In  older  stipes  the  pith  web  is  decidedly  solid,  in  marked  con- 
tradistinction to  this  tissue  in  young  material.  As  the  stipe 
matures  the  interstices  between  the  elements  of  the  pith  web 
become  obliterated  by  the  repeated  branching  and  interlacing  of 
the  filaments.  The  chlorophyll  also  disappears  and  the  cells 
become  filled  with  densely  granular  contents.  There  remain, 
however,  in  even  the  oldest  pith,  numerous  interstitial  passages 
which,  in  cross  section  or  longitudinal  section,  present  much  the 
same  appearance  and  are,  perhaps,  what  were  mistaken  for 
mucilage  ducts  by  Ruprecht. 

A  cross  section  of  the  mature  stipe  shows  then  the  following 
characters.  At  the  center  is  the  solidly  interwoven  tissue  of 
the  pith  web.  This  is  surrounded  by  the  clearly  marked, 
sclerenchymatous  tissue  of  the  primary  cortex  passing  insensi- 
bly into  the  tracheid-like  tissue  of  the  secondary  cortex  which 
is  arranged  in  concentric  rings,  resulting  from  the  succession 
of  elements  with  larger  cell-lumina  in  apposition  upon  those 


MacMillan:  OBSERVATIONS  ON  PTERYGOPHORA.  737 

with  smaller.  Outside  of  the  rings  will  be  found  the  cambial 
zone  in  which  regular  divisions  take  place  in  all  three  planes  of 
space.  Exterior  to  the  cambial  zone  lies  a  thin  outer  cortex 
composed  of  cells  very  much  smaller  than  those  of  the  inner 
area,  provided  with  thick  walls  and  constituting  a  kind  of  bark 
for  the  trunk.  In  some  material  the  general  cambial  zone  can 
be  very  distinctly  seen,  ten  or  twelve  cells  in  thickness  and 
separated  from  the  epidermis  by  twenty  or  more  layers.  Not 
infrequently  the  cells  at  the  periphery  of  a  ring  of  growth  have 
more  densely  granular  contents  than  those  of  the  general  sec- 
ondary cortex  tissue.  Thus,  occasionally  in  the  stipe  there  may 
arise  the  anatomical  conditions  which  seem  to  be  more  normally 
characteristic  of  the  holdfast.  The  photographs  of  different 
cross  and  longitudinal  sections  which  are  presented  will  serve 
to  make  these  points  clear  where  the  description  is  necessarily 
difficult  to  follow. 

The  lamina. — As  before  stated  only  the  central  lamina  is  pro- 
vided with  a  midrib,  the  pinnae  being  quite  devoid  of  such  a 
structure.  The  midrib  of  the  central  lamina  arises  through  an 
hypertrophy  of  the  cortical  tissue,  in  which  th  ,  pith-plate  does 
not  seem  to  partake.  The  general  structure  of  the  lamina  as 
seen  in  cross  section  does  not  present  many  peculiar  features, 
but  is  much  like  that  already  described  for  other  genera.  There 
is  on  each  surface  an  isomorphic  epidermis  composed  of  small 
quadrate  chlorophyll-containing  cells,  and  these  merge  insensi- 
bly into  the  subepidermal  tissue,  which  in  some  instances  is 
two  or  more  layers  in  depth.  The  cells  then  become  much 
larger  in  diameter  and  the  contents  less  dense.  Among  these 
cortical  cells  occasional  very  large  polysaccharid  idioblasts  are 
found,  and  in  the  chocolate-colored  pinnae  of  plant  "  C,"  these 
cells  are  very  numerous  and  densely  packed  with  spherical 
bodies,  doubtless  belonging  to  the  category  of  reserve  carbohy- 
drates. Owing  to  the  nutritious  character  of  such  pinnae,  they 
are  very  commonly  perforated  by  animals,  sometimes  giving  a 
colander  appearance  like  that  of  Agarum,  and  covered  with 
epiphytic  and  endophytic  vegetation,  a  further  study  of  which 
should  be  made.  These  reservoir  cells  may  perhaps  be  packed 
with  food  materials  previous  to  the  production  of  sori  and  the 
polysaccharids  utilized  in  the  elaboration  of  the  sporangia  and 
paraphyses.  In  any  event  they  seem  to  be  emptied  of  their 
contents  underneath  most  of  the  soral  areas  that  I  have  exam- 


738  MINNESOTA    BOTANICAL    STUDIES. 

ined.  Within  the  layer  of  idioblasts  the  cortical  cells  become 
smaller  on  each  side  of  the  pith  plate.  This  latter  is  composed 
of  anastomosing  filaments  with  occasional  trumpet-hyphas  and 
does  not  differ  particularly  from  the  same  tissue  as  displayed  in 
the  stipe.  The  cortex  of  the  mid-lamina  and  of  the  pinnae 
seems  to  show  some  fairly  constant  differences.  In  the  pinnae 
the  cells  are  often  uniformly  larger  and  the  layer  of  idioblasts 
is  frequently  almost  continuous. 

The  sorus. — I  do  not  find  anywhere  in  the  literature  of  Ptery- 
gophora  an  account  of  its  fruiting  area.  The  first  fruiting  ma- 
terial that  reached  me  was  collected  near  Port  Renfrew  by  Miss 
Josephine  E.  Tilden  during  December,  1901.  Shortly  after- 
wards Professor  Setchell  kindly  sent  me  some  dried  pinnae  col- 
lected at  Whidby  Island,  Washington,  by  Mr.  N.  L.  Gardner, 
and  said  to  display  sori.  My  observations  are  made  from  the 
fresh  material  collected  by  Miss  Tilden.  The  soral  patches 
occur  upon  the  lateral  pinnae  and  not  upon  the  mid-lamina.  In 
this  respect  they  remind  one  of  the  eoral  distribution  of  Alaria ; 
but  while  in  Alaria  the  entire  pinna,  except  a  very  small  mar- 
ginal region,  is  soriferous,  in  Pterygophora  the  sori  form  some- 
what irregular  patches  upon  both  surfaces  of  the  pinna  and 
much  of  its  area  fails  to  develop  them.  In  this  respect  Ptery- 
gophora  approaches  more  nearly  to  Laminaria. 

The  sori  are  composed  of  numerous  elongated,  saccate  goni- 
dangia,  each  bearing  from  fifty  to  two  hundred  spherical  go- 
nidia.  The  gonidangial  surface  is  surmounted  by  the  inter- 
mingled paraphyses,  averaging  half  as  long  again  as  the 
gonidangia  and  their  clavate  distal  ends  capped  with  cuticular 
masses,  suggesting  those  already  described  for  Lessonia  and 
indicating  that  in  this  plant,  as  in  Lessonia,  the  cuticular  layer 
does  not  separate  in  a  plate,  as  described  for  Laminaria  by 
Thuret  and  as  known  to  occur  also  in  Nereocystis,  but  divides 
into  individual  paraphysal  calyptra.  Some  gonidangia  meas- 
ured indicated  an  average  length  of  50  mic.  and  diameter  of 
10  mic.  The  gonidia  are  nearly  2  mic.  in  diameter  and  in  a 
double-stained  preparation,  made  for  me  by  Mr.  H.  L.  Lyon, 
take  Delafield's  haematoxylin  while  the  paraphyses  and  cutic- 
ular substance  take  the  safranin.  The  gonidangia  and  para- 
physes stand  upon  a  floor-layer  such  as  is  found  in  all  Laminaria- 
ceous  sori,  and  their  development  does  not  differ  from  that 
which  has  already  been  described  for  other  genera. 


MacMillan:  OBSERVATIONS  ON  PTERYGOPHORA.  739 

I  desire  to  express  my  thanks  to  Miss  Josephine  E.  Tilden 
for  the  winter  fruiting  material  which  she  kindly  collected  for 
me  and  for  the  series  of  slides  from  which  most  of  my  descrip- 
tion has  been  made,  and  to  Mr.  H.  L.  Lyon  for  the  photo-mi- 
crographs reproduced  in  plate  LXI  and  prepared  by  him  at 
my  request. 

SUMMARY. 

1.  Ptcrygophora    calif ornica    grows    to   a  larger    size  than 
generally  known.     Specimens  ten  feet  in  length  with  trunks 
three  inches  in  diameter  have  been  seen. 

2.  As  displayed  in  the  Straits  of  Fuca,  Pterygophora  is  a 
surge  plant,  growing  below  the  zone  of  Lessonia  and  above 
that  of  Nercocystis. 

3.  Pterygophora  may  be  classified  either  in  the  Laminariese 
or  the  Alariideae.     Its  characters  are  in  many  respects  inter- 
mediate between  these  tribes. 

4.  The  holdfast  shows  distinct  rings  of  growth  and  these  in 
most  instances  arise,  not  through  morphological  differences  be- 
tween adjacent  cell-layers,  but  through  differences  in  the  cell 
contents.     The  substances,  which  produced  in  greater  or  less 
amount  give  the  ringed  appearance,  are  regarded  as  polysac- 
charids  allied  to  mucine,  as  described  by  Koch. 

5.  The  stipe  is  devoid  of  the  mucilage  ducts  of  Ruprecht  and 
shows  distinct  rings  of  growth,  due  in  most  instances,  to  the 
juxtaposition  of  a  layer  of  cells  with  larger  lumina,  upon  a  layer 
with  smaller.     In  some  cases  the  ringed  appearance  of  the  stipe 
seemed  to  be  due  to  the  same  condition  described  for  the  holdfast. 

6.  In  the  cortex  of  the  lamina  large  polysaccharid  idioblasts 
are  abundantly  developed.     These  are  most  numerous  in  the 
pinnas  and  are  often  exhausted  of  their  contents  during  the  proc- 
ess of  soral  formation. 

7.  The  sori  are  distributed  in  irregular  patches  toward  the 
base  of  the  pinna  and  in  the  disposition  of  the  cuticular  caps 
upon  the  paraphyses  suggest  Lessonia.     The  plant  fruits  in  the 
latitude  of  Port  Renfrew  during  the  month  of  December. 

DESCRIPTION  OF  PLATE  LVII. 

Young  plants  of  Pterygophora,  about  one-half  natural  size. 

1.  Plant  "A"  of  text. 

2,  3,  4.   Somewhat  older  plants;   in  4  the  midrib  is  just  beginning 
to  appear  in  the  base  of  the  lamina. 


740  MINNESOTA    BOTANICAL    STUDIES. 

5.  Plant  "  B  "  of  text;  the  midrib  is  more  distinct  and  the  two  op- 
posite growing  points  of  the  first  pair  of  pinnae  may  be  seen  just  be- 
low the  junction  of  lamina  with  stipe. 

PLATE  LVIII. 

Plant  "  C  "  of  text.  The  plant  is  shown  a  little  more  than  one- 
third  the  natural  size.  It  presents  the  loosely  arranged  pinnae  charac- 
teristic of  young  plants  and  as  shown  in  Ruprecht's  plate.  The  lower 
pinnae  are  perforated  by  marine  animals  and  are  the  ones  in  which  the 
reserve  material  is  the  most  abundant.  The  mid-lamina  alone  shows 
a  midrib. 

PLATE  LIX. 

Plant  "  D  "  of  text.  The  size  is  indicated  by  the  hat  placed  in  the 
field  of  view.  It  is  about  one-tenth  natural  size  and  was  photographed 
on  the  shore  a  few  moments  after  collection.  It  is  somewhat  fore- 
shortened, being  between  three  and  four  meters  in  length.  The  cen- 
tral lamina  is  shown  in  line  with  the  stipe. 

PLATE  LX. 

Pinna-region  of  full-grown  specimen,  one-half  natural  size.  The 
very  much  crowded  position  of  pinnae  in  old  plants  is  indicated  in  this 
plate.  Below  the  pinnae  may  be  seen  scars  left  by  pinnae  of  previous 
seasons. 

PLATE  LXI. 

Cross  sections  through  the  stipe,  natural  size.  The  lower  figure  is 
taken  near  the  base  of  the  stipe,  while  the  upper  is  cut  just  below  the 
region  of  pinnae.  The  great  difference  in  the  extent  of  the  pith  and 
in  its  shape,  in  the  two  sections,  is  noticeable.  The  characteristic 
rings  of  growth  are  apparent  in  both  figures. 

Plates  LVII.  to  LXI.  are  from  photographs  made  under  the  direc- 
tion of  the  author,  by  Mr.  C.  J.  Hibbard,  Photographer  of  the  De- 
partment of  Botany  in  the  University  of  Minnesota. 

PLATE  LXII. 

The  anatomical  detail  of  Pterygophora. 

1.  Cross  section  through  secondary  cortex  of  stipe,  showing  rings 
of  growth  (  x  50). 

2.  Cross  section  through  stipe,  showing  secondary  cortex  above  and 
primary  cortex  below  (x  50). 

3.  Longitudinal  section  through  secondary  cortex  showing  flatten- 
ing of  cells  towards  the  right.      This  would  appear  as  a   ring   in   the 
cross  section  (  x  50). 


MacMillan:  OBSERVATIONS  ON  PTERYGOPHORA.  741 

4.  Tangential  section  through  secondary  cortex,  showing  the  some- 
what fusiform  outline  of  the  cells  in  this  section  (x  50). 

5.  Longitudinal  section  through  region  of  Fig.  3.     On  the  left  is 
seen  the  tissue  of  the  secondary   cortex,  in  the  middle  the  flattened 
cells  of  the  transition  zone,  and  on  the  right  the  cells  of   the  primary 
cortex.     This  first  ring  of   growth,  caused  by  the  superposition  of 
secondary    upon   primary  tissue,    is   the    most   prominent   of    all    the 
growth  rings  in  the  stipe  (  X  50). 

6.  A  portion  of  the  secondary  cortex  tissue  shown  in  Fig.  5,  mag- 
nified to  demonstrate    the  pits  in  the  vertical  walls  of  the  tracheid- 
likc  elements.     To  the  right  the  cells  become  compressed,  passing 
into  the  transition  zone  (  X  250). 

7.  A  portion  of  the  primary  cortex  tissue  shown  in  Fig.  5,  mag- 
nified   to    demonstrate    the    different    arrangement   of    primary    and 
secondary  cortical  cells.     The  secondary  stand   in   long  rows  at  the 
same  level.      This  is  not  true  of  the  primary  (  X  250). 

8.  Cross  section  through  one  of  the   rings   shown  in  Fig.  i   and 
magnified  to  demonstrate  the  flattening  of  cells  in  the  region  of  the 
ring  and  their  denser  contents.     The  appearance  of  small  cells  inter- 
polated between  the  larger  ones  can  be  understood  by  referring  to  Fig. 
4  (X   250). 

9.  Cross  section  through  pith  web  of  stipe,  showing  anastomosing 
filaments  and  trumpet  hyphaB  embedded   in  a  gelatinous  matrix  (x 
250). 

10.  Cross  section  through  haptere.     The  cells  of  the  outer  cortical 
region  are  crowded  with  contents,  and  do   not  differentiate  clearly. 
Near  the  middle  of    the   section  are  seen  two  of    the  characteristic 
growth-rings  of  the  haptere  (x  50). 

1 1.  Longitudinal  section  through  haptere,  showing  the  development 
of  secondary  tissues,  the  cells  of  which  are  shorter  upon  the  primary 
tissue  towards  the  bottom  of  the  figure.     Below  the  black  outer  cortex 
a  growth-ring  in  section  may  be  seen  (  x  60). 

12.  Section   through   sorus,   showing    floor-cells,   gonidangia  with 
gonidia  and  paraphyses  capped  with  cuticular  knobs  characteristic  of 
this  genus  and  of  Lessonia  (  x  320). 

All  the  figures  in  Plate  LXII.  are  from  original  photomicrographs 
by  Mr.  H.  L.  Lyon. 


VOL.  II.  MINNESOTA  BOTANICAL  STUDIES.  PART  VI. 


PLATE  LVII. 


MINNESOTA  BOTANICAL  STUDIES. 


PART  VI. 


VOL.  II. 


MINNESOTA  BOTANICAL  STUDIES 


PART  VI. 


PLATE  LIX. 


VOL.  II. 


MINNESOTA  BOTANICAL  STUDIES. 


PART  VI. 


PLATE  LX. 


HELIOTYPE    CO.,    BOSTON 


VOL.  II. 


MINNESOTA  BOTANICAL  STUDIES. 


PART  VI. 


PLATE  LXI. 


MINNESOTA  Bcr 


r 


m 

.•.•<•- 

>UJU/jAi 


L  STUDIES. 


PART  VI. 


-   /f 


10 


INDEX   OF   PLANT   NAMES.* 


Abies  balsamea  (L. )  Mill.,  225,  559 
Abutilon  abutilon  (L. )  Rusby,  397 
Acalypha  virginica  L.,  395 
Acer,  88,  564,  566 

glabruin  Torr.,  75,  77 

negundo  Linn.,  77,  84,  85,  362,  396, 
586 

nigrum  Michx.,  362,  396 

saccharum  Marsh.,  586 

saccharinum  Linn.,  77,  85,  358,  396 

spicatum  Lam.,  225,  272,  366,  396 
Aceracese,  77,  396,  586 
Acerates  viridiflora  (Raf. )  Eaton,  367, 

402,  558 

Achillea  millefolium  L.,  414,  598 
Acnida   tamariscina    (Nutt. )    Wood., 

358,  385 
Acolium   tigillare    (Ach.)    DN.,*  306, 

673,  706 
Aconitum,  347 

abbreviatum  Langsd.,  347 

acuminatum  Reichenbach,  347 

acutum,  351 

cegophonum,  349 

album  Ait.,  350 

alien  urn,  349 

alpinum  Mill.,  350 

altigaleatum  Hayne,  350 

altissimum  Mill.,  349 

atnbiguum,  351 

amcenum,  351 

Anthora  Linn.,  352 

Anthorideum  DC.,  352 

Anthorum  St.  Lag.,  352 

arctophonum,  349 

arcuatum  Maxim.,  347 

australe,  349 

austriacum  Tratt.,  346 

autuntnale  Lind.,  347 

autumnale  Reichb.,  352 

barbatum  Patr.,  349 

Bernharaianum,  351 


Aconitum 

bicolor  Schult.,  346 
Braunii,  351 
Breiteriatium,  346 
bulbiferum  Howell,  348,  350 
Burnhaidianum  Wallr.,  350 
Californicum  Hort.,  347 
callibotryon,  351 
camarum  Schleich.,  347 
cammarum  Jacq.  ,  350 
cammarum  Linn.,  346 
Candollei,  352 
cernutim  Baumg.,  346 
cernuum  Wulf.,  347 


dusiamtm,  351 
Columbianum  Nutt.,  347 
commutatum,  351 
cordatum  Royle,  348 
cynodonum,  349 
decorum,  346 
delphinifolium  Hort.,  349 
delphinifolium  DC.,  351 

var.  ramosum  n.  var.,  351 
dissectum  Don.,  351 
disseclum  Tausch.,  349 
eriostemum  DC.,  346 
eulophum,  352 
exaltatum,  346 
ex  eel  sum  Turcz.,  349 
firmum,  351 
Fischeri  Rchb.,  347 
flexuosum  Presl.,  350 
formosum,  351 
Funkianum,  351 
galectonum,  347 
galei-iflorum  Stokes,  349 
gibbiferum,  347 
glabrum  DC.,  350 
Gmelini,  349 
gracile,  350 
hamatum,  346,  350 


The  index  was  prepared  by  Miss  Josephine  E.  Tilden. 

743 


744 


INDEX. 


Aconitum 

hebegynum  DC.,  347 
heterophyllum  Wall.,  348 
hians,  351 
hispidum  DC.,  349 
Hoppeanum,  351 
hortense,  346 
Hosteanum  Schur.,  349 
humile  Salisb.,  347 
il/initum,  350 
intermedium  DC.,  346 
intermedium  Gaud.,  350 
intermedium  Host.,  349 
italic  inn  Tratt.,  350 
Jarquini,  352 
Jacquinianum  Host.,  349 
Japonicum  Hort.,  350 
Japonicum  Thunb.,  348 
Japonicum     Decne. — var.     cceru- 

leum  Hort.,  348 
Kamtschaticum  Pall.,  347 
Koehleri,  351 
Lahanskyi  Rchb.,  347 
laciniosum  Schleich.,  350 
leetum,  351 

tevigatum  Schleich.,  350 
lagoctonum,  349 
Lamarckii,  349 
lasiocarpum,  350 
laxiflorum  Schl.,  351 
laxum,  351 

leucanthemum  Wender,  350 
loreale  Sen,  349 
luparia,  349 
lupicida,  349 
luridum  Salisb.,  350 
Lycoctonum  Lind.,  349 
macranthum,  350 
maximum  Pall.,  347 
meloctonum,  349 
Mielichhoferi,  351 
ntixlum,  350 
moldavicurn,  349 
molle,  347 

monavense  Schmidt,  350 
myoctonum,  350 
Napellus  S.  G.  Gmel.,  349 
Napollus  Linn.,  350 
Napellus  Thunb.,  347 
Napellus  var.  delphinifolitim  Sc- 
ringe, 351 


Aconitum 

napelloides  Sw.,  351 
nasutum  Hook.,  347 
nasntum  Fisch.,  350 
Neapolitanum  Tenore,  349 
nemerosum  Bieb.,  352 
iieomontanum  Baumg. ,  347 
neomontanum  Willd.,  346 
neubergense  DC.,  351 
Noveboracense  Gray,  347 
ochranlhum  C.  A.  Mey.,  349 
ochroleucum  Hort.,  349 
ochroleucum  Salisb.,  352 
ochroleucum  Willd.,  349 
oligocarpum,  351 
Ottonianum,  346 
Pallasii,  352 
pal/idum,  350 
palmatifidum,  346 
paniculatum  Lam.,  347 
paradoxum,  351 
parviftorum  Hoppe  &  Horn,  347 
pauciflorum  Host.,  349 
perniciosum,  350 
Phthora,  350 
plexicau/e,  347 
pyratnidale  Mill.,  350 
Pyrenaicum  Linn.,  349 
Pyrenaicum  Pall.,  352 
ramosum  A.  Nels.,  351 
ranunculifolium,  350 
reclinatum  Gray,  352 
tectum  Bernh.,  350 
reflexum,  347 
rhynchanthum,  350 
rigidum,  351 
rostratum  Bernh.,  350 
rubicund  it  in  Fisch.,  349 
scandens  Muhl.,  348 
semigaleatum,  351 
septentrionale  Koelle.,  349 
sinense  Sieb.  &  Zucc.,  347 
speciosum  Otto,  346 
Sprengelii  Rua,  346 
squarrowm  Linn.,  349 
Storkianum,  346 
Storkianum  Rchb.,  347 
strictissimum,  350 
strictum  Bernh.,  351 
strictum  Willd.,  350 
tauricum  Wulf.,  351 


INDEX. 


745 


Aconitum 

taxicarium  Salisb.,  349 

taxi  cum,  347 

tenuifolium,  351 

thelyphonum,  350 

theriophonum,  350 

tragoctonum,  350 

Transilvanicum  Lerch.,  349 

triste  Fisch.,  349 

tuberosum  Pair.,  352 

umbracticolum  Schur.,  349 

uncinatum  Hort.,  350 

uncinatura  Linn.,  348 

varicgatum  Hook.,  348 

variegatum  Linn. — var.  album  n. 
var.,  350 

venustum,  351 

versicolor,  346 

virgatum,  351 

volubile  Moench.,  350 

volubile  Muhl.,  348 

vu/paria,  350 

Wilematianum  Delarb.,  347 

zooctonum,  350 
Acorus  calamus  L.,  543,  576 
Actsea  alba  ( L. )  Mill. ,  362,  387,  555,  581 

rubra  (Ait.)  Willd.,  362,  387 
Adiantum  pedatum  L.,  362,  371 
Adicea  pumila  (L. )  Raf.,  384 
Adopogon    virginicum    (L. )    Kuntze, 

362,  409,  556 
Adoxa  moschatellina  IyM  362,  366,  408, 

544 
Adoxaceae,  408 

,  537,  554,  631 

actseae  Opiz.,  555 

album  Clint.,  541 

asterum  Schwein.,  557 

cimicifugatum  Schwein.,  555 

clematidis  DC.,  555 

compositarum  Mart.,  556 

var.  erigerontis  Wint.,  556 
var.  eupatorii  (Schw. )  Burrill, 

557 

var.  helianthi  Burrill,  557 
var.  lactucae  Burrill,  557 
var.  liatrii  Webber,  557 
var.  prenanthis   (P.)    Wallr., 

556 

compositarum,  557 
convallariae  Schum.,  554 


elatinum,  559 

fraxini  Schwein.,  559 

fumariacearum  Kell.  &  Swingle, 
555 

geranii  DC.,  556 

grossulariae  Pers.,  558 

grossularics  Schum.,  558 

hydnoideum  B.  &  C.,  558 

hydrophylli  Peck,  559 

impatientis  Schwein.,  556 

iridis  Ger.,  554 

jacobeae  Grev.,  556 

jamesianum  Peck,  558 

lupini  Peck,  554 

lycopi  Ger.,  546 

lysimachiae  (Schl. )  Wallr.,  558 

Mariae-Wilsoni  Peck,  632,  639 

orobi  Pers.,  554 

pammelii  Trel.,  559 

peckii  DeToui,  559 

pedatatum    (Schw.)    Arthur    and 
Holway,  632 

Petersii  B.  &  C.,  632 

phrymae  Halst.,  559 

porosum  Pk.,  541 

pulcherrimum  Rav.,  633 

punctatum  Pers.,  555 

pustulatum  Curt.,  558 

rauunculacearum  DC.,  555 

ranunculi  Schwein,  555 

rubellum  Gmel.,  547 

senecionis  Desmaz,  556 

thalidri  Grev.,  555 

thalictri-flavi  (DC.)  Wint.,  555 

urticas  Schwein.,  546 

uvularias  Schwein.,  554 

verbenae  Speg.,  556 

violce  Schum.,  633 

violarum  DC.,  633 
Agarum,  737 

Agastache   anethiodora  (Nutt. )    Brit- 
ton,  591 

scrophulariae folia   (Willd.) 

Kuntze,  362,  404 

Agoseris   glauca   (Pursh^  Greene,  594 
Agropyron  repens  (  L. )  Beauv.,  574 

tenerum  Vasey,  575 
Agrimonia  hirsuta  (Muhl.)  Bicknell, 

362,  391.  56o,  584 
Agrostemma  githago  L.,  581 


746 


INDEX. 


Agrostis  alba  L..  573 

Ailanthus  glandulosa  Desf.,  76,  85,  87, 

112,  130,  131,  132,  133 
Aizoaceae,  386 
Alaria,  561,  723,  728,  730,  732 

angusta,  562 

curtipes  Saunders,  561 

esculenta  (L- )  Grev.,  725 

praelonga,  561 
Alectoria  jubata  (L. )  Tuck.,  219,  243, 

659,  677,  684 
var.  chalybeiformis  Ach.,  244, 

675,  677,  684 
var.  implexa  Fr.,  244 

ssepincola  (Ehrh.)  Ach.,  228 
Alisma  plantago,  144 

plantago-aquatica     L.,    356,    372, 

373,  626 

Alismaceae,  373,  572 
Alismineae,  651 
Allionia  diffusa  Heller,  33 

hirsuta  Pursh,  581 

linearis  Pursh,  368,  386 

nyctaginea  Michx.,  385 
Allium  stellatum  Ker.,  577 
Alnus,  430,  565 

alnobetula  (Ehrh.)  K.  Koch,  578 

incana  (L,.)  Willd.,  382,  427 
Alopecurus  geniculatus  L. ,  573 
Alsine  media  L.,  581 
Amaranthaceae,  385,  580 
Amaranthus  blitoides  S.,  385,  580 

graecizans  L-,  580 

retroflexus  L.,  385,  580 
Amaryllidaceae,  379 
Amblystegium  adnatutn  L.  and  J.,  50 

serpens  Sch.,  50 
Ambrosia,  551 

artemisisefolia  L-,  409,  425,  594,  625 

psilostachya  DC.,  425,  594 

trifida  L.,  409,  425,  55 r,  594,  626 
Ambrosiaceae,  409,  594 
Amelanchier  alnifolia  Nutt.,  391 

botryapium  (L,.  f.)  DC.,  391 

canadensis  (L. )   Medic.,  362,  391 
Amorpha,  84,  544,  552 

canesceus  Pursh,   20,    21,  22,    23, 
393,  552 

fruticosa  Linn.,  74,  87,   107,   131, 

132,  133,  393,  553,  584 
nana  Nutt.,  74,  584 


Amphiroa  calif ornica  Dene.,  701,  703 

cretacea  Endl.f.tasmanica  (Lond.) 
Yendo,  700,  701,  702,  703 

epiphlegnoides  J.  Ag.,  701,  703 

mallardiae  Harv.,  706 

tasmanica  Lond.,  701 

tuberculosa  Endl.,  700,  701,  702 

wardii  Harv.,  706 

Amphora  ovalis  (Breb.)  Kg.,  613,  614 
Amphoridium  lapponicum  Sch.,  41 
Anacystis    marginata    Menegh.,    613, 

614 
Anabaena,  6n,  614 

azollae  Strasb.,  27 

cycadearum  Reinke,  27 

flos-aquae  (Lyngb. )  Breb.,  27,  612 
Anacardiaceae,  76,  114,  395,  586 
Andropogon  furcatus  Muhl.,  374,  572 

scoparius  Michx.,  22,  572 
Anemone  canadensis  L.,  387,  555,  581 

cylindrica  A.  Gray,  581 

nudicaulis  Gray,  491 

quinquefolia  L.,  362,  387,  555, 

virginiana  L.,  387,427,  581 
Angelica  atropurpurea  L.,  361,  399 
Anogra  albicaulis  (Pursh)  Britton,  427 

pallida  (Lindl. )  Britton,  587 
Anomodon  attenuatus  Hartm.,  48 

minor  (P.  Beauv. )  Fiirn.,  48 

rostratus  Sch.,  48 
Anomodonta,  36 
Antennaria  campestris,  20,  21,  24 

plantaginifolia  (L.)  Richards,  412 
Anthemis  cotula  L.,  414,  622 
Anthoceros,  195 

Apios  apios  (L-)  MacM.,  362,  394,  554 
Aplozia  autumnalis  (DC. )  Heeg,  193 
Apocynaceae,  401,  589 
Apocynum  androsEemifolium  L.,   362, 
401,  589 

cannabinum  L.,  358,  401 

cannabinum  glaberrimum  DC., 401 
Aquilegia,  331 

agger icola  Jord.,  338 

alpina  Linn.,  333,  343 

alpina  var.  superba  Hort.,  343 

arctica  Loud.  Hort.,  340 

atrata  Koch.,  337 

alropurpurea  Miquel.,  336 
•    atroputpurea  Willd.,  336 

aurea  Junka.,  341 


INDEX. 


747 


Aquilegia 

Bernardi  Green  &  Godr.,  338 

bicolor  Ehrh. ,  339 

var.  flore-pleno  Hort.,  339 

blanda  Lew.,  339 

brachyceras  Turcz.,  337 

brevistyla  Hook.,  334 

Buergeriana,  333 

Buergeriana  Sieb.   &   Zucc.,   var. 
ecalcarata  Davis,  336 

ccerulea  James,  332,  333,  342 
var.  albiflora  Gray,  342 
var.  alpina  A.  Nelson,  342 
var.  calcarea  Jones,  342 
\wc.flavescens  Lawson,  335 
var.  flore-pleno  Hort.,  343 
var.  hybrida  Hort.,  342 

Californica  Lindl.,  340 

var.  hybrida  Hort.,  340 

Canadensis  Linn.,  335,  387 
var.  aurea  Roezl.,  341 
var.  depauperata  Fink,  335 
var.  flaviflora  Britton,  335 
\&r.formosa  Wats.,  340 
var.  nana  Hort.,  336 

Caucasica  Ledeb.,  339 

chrysantha  Gray,  333,  341 

var.  alba-plena  Hort.,  341 

var.  aurea  Davis,  341 

var.    grandiflora-alba    Hort., 

34i 

var.  Jacschkani  Hort.,  341 
var.  nana  Hort.,  341 

col  Una  Jord.,  338 

concolor  Fisch.,  338 

corniculata  Vill.,  337 

cornuta  Gilib.,  337 

Corsica  Solier.,  338 

dahurica  Patr.,  336 

dumeticola  Jord.,  338 

depauperata  Jones,  335 

ecalcarata  Eastwood,  337 

ecalcarata  Hort.,  338 

elata  Ledeb.,  337 

elegans  Pope,  338 

elfgans  Salisb.,  335 

elegantula  Greene,  334 

eximia  Van  Houtte,  340 

flabellata  Sieb.  &  Zucc.,  333,  337 

var.  nana-alba  Hort.,  337 
flavescens  Wats.,  335 


Aquilegia 

flaviflora  Tenney,  335 

flavescens,  335 

var.  hybrida  Hort.,  340 
var.  nana-alba  Hort.,  340 
var.  rubra-pleno  Hort.,  340 
var.  truncata  Baker,  340 

formosa  Fisch.,  333,  340 

var.  desertorunt  Jones,  340 

Gamietiana  Sweet,  339 

glandulosa  Fisch.,  332,  333,  343 
var.  jucunda  Fisch.    &  Lall., 

343 

glandulosa  Miq.,  337 
glaucescens  Baker,  338 
glaucophylla  Steud.,  338 
Haenkeana  Koch.,  338 
inversa  Mill.,  337 
Jonesii  Perry,  334 
Karelini  Baker,  338 
lactiflora  Kar.  &  Kir.,  333 
Laramiensis  A.  Nelson,  334 
leptocera  Nutt.,  342 
leptoceras  Fisch.   &  Meyer,  333, 

337 

var.  chrysantha  Hook.,  341 
\ar.Jlava  Gray,  341 
var.  lutea  Hort.,  341 

longissima  Gray,  333,  341 

macrantha  Hook.  &  Arn.,  342 

Mexicana  Hook.,  339 

micrantha  Eastwood,  333,  336 

montana  Sternb.,  343 

nigricans  Baumg.,  338 

Olympica  Boies,  339 

oxyscpala  Traut.  &  Meyer,  333 

paraplesia  Schur.,  338 

platysepala  Reichb.,  338 

plena  Hort.,  338 

pracox]m&.,  337,  338 

pubescens  Coville,  333,  341 

Reuteriana  Reichb.,  343 

saximontana,  P.  A.  Rydberg,  335 

Sibirica  Lam.,  333,  339 

var.  flore-pleno  Hort.,  339 
var.  spectabilis  Baker,  339 

sibirica  Don,  338 

silvestris  Neck,  337 

Skinneri  Hook.,  333,  339 

var.  flore-pleno  Hort.,  340 
var.  hybrida  Hort.,  341 


748 


INDEX. 


Aquilegia 

speciosa  DC.,  339 

speclabilis  Lem.,  339 

stellata  Hort.,  337 

Sternbergii  Reichb.,  343 

Stuarti  Hort.,  343 

subalpina,  338 

subscaposa  Borhas,  338 

sylvesfris  Schur.,  338 

Ttanssilvanica  Schur.,  338 

truncata  Fisch.,  340 

variegata  Moench.,  335 

versicolor  Salisb.,  337 

viridi  flora     Pallas     var.     atropur- 

purea,  336 

vulgaris  Linn.,  332,  337 
var.  alba  Hort.,  338 
var.  atroviolacea  Hort.,  339 
var.  brevistyla  Gray,  334 
var.  hybrida  Sims,  339 
var.  flora-piano  Hort.,  338 
var.  folio-aureis  Hort.,  339 
var.  nivea  Bautng.,  338 
var.  Olympica  Baker,  339 
var.  Olympica,  343 
var.  vervteneana  Hort.,  339 
vulgaris  Richards.,  334 
vulgaris  Thunb. ,  337 
Wittmanniana  Stev.,  339 
Arabis  canadensis  L.,  389 
glabra  (L. )  Bernh.,  389 
laevigata  (Muhl. )  Poir.,  389 
Araceae,  377,  576 
Aragallus  involutus  A.  Nels.,  427 
Aralia  nudicaulis  L-,  362,  399,  554,  588 

racemosa  L.,  362,  398 
Araliacese,  398,  588 
Archispermae,  372 
Arcteranthis  Greene,  502 
Cooleyae  Greene,  502 
Arctium  lappa  I/.,  598 
minus  Schk.,  415 
Arctostaphylos  uva-ursi  (L. )  Spreng., 

368,  400,  588 
Argentina    anserina  .(L.)    Rydberg, 

583 
Arisaema,  55,  540 

dracontium,  358,  377 

triphyllum   (L. )  Torr.,  62,  63,  65, 

362,  377,  542,  576 
Aristolochiaceae,  384 


Artemisia  absinthium  L-,  598 

biennis  Willd.,  598 

caudata  Michx.,  598 

dracunculoides  Pursh,  414,  545,  598 

frigida  Willd.,  598 

gnaphalodes   Nutt.,    22,    23,    414, 
598,  627 

serrata  Nutt.,  414 
Arthonia  sp.,  281,  326,  705 

dispersa  (Schrad. )  Nyl.,  225,  272, 
304,  326,  705 

lecideella  Nyl.,  303,  304,  325,  659, 

6?i,  705 

patellulata  Nyl.,  229,  273,  705 

punctiformis  Ach.,  273,  305,  326 

radiata  (Pers. )  Th.   Fr.,  272,  303, 

326,  705 
Arthrocardia ?  frondescens   (Post,    et 

Rupr. )  Aresch.,  704 
Arthrodesmus  incrassatus  Lagerh.  var. 

cycladatus  Lagerh.,  612,  614 
Asarum  canadense  L.,  362,  384 
Asclepiadaceas,  401,  589 
Asclepias exaltata  (L. )  Muhl.,  362,  401 

incarnata  L.,  357,   362,  401,    589, 
626 

obtusifolia  Michx.,  368,  401 

speciosa  Torr.,  590 

sullivantii,  626 

syriaca  L.,  362,  401,  558,  589 

tuberosa  L.,  401,  558 

verticillata  L-,  367,  402 
Asplenium    acrostichoides    Sw.,    362, 
366,  370 

angustifolium  Michx.,  368,  370 

filix-fcemina  (L. )  Bernh.,  362,  366, 

371 
Aster,  550 

divaricatus  L.,  539 
drummondii  Lindl.,  411 
dumosus,  627 
Isevis  L.,  411,  596 
lindleyanus  T.  &  G.,  620 
macrophyllus  L-,  539,  550 
multiflorus  Ait.,  20,  21,  23,  596 
novae-anglise  L.,  361,  411,  595 
paniculatus  Lam.,  412,  596 
patens  Ait.,  411 
prenanthoides  Muhl.,  361,  411 
ptarmicoides  (Nees)  T.  &  G.,4ii, 

596 


INDEX. 


749 


Aster 

puniceus  L,.,  361,  411,  596 

var.  lucidulus  A.  Gray,  425 
sagittifolius  Willd.,   23,   361,  411, 

550,  557.  595 

salicifolius  Lam.,  411,  596 
sericcus  Vent.,  367,  411,  596 

Asterionella  formosa  Hass.,  611,  613, 
614 

Astragalus  canadensis  L.,  427 
carolinianus  L.,  393,  585 
flexuosus  (Hook.)  Doug.,  192 

Atragene  americana  Sims,  387 

Atricbum  uudulatum  P.  B.,  43 

Atriplex  patula  L.,  569,  580 

Aut-eupuccinia,  543,  544 

Aut-euromyces,  540 

Avena  sativa  L.,  548 

Azolla  caroliniana  Willd.,  27,  355,  368, 
371 

Baeomyces,  236,  662 

aeruginosus     (Scop.)      DC.,    233, 
266 

byssoidcs  (L. )  Schaer.,  227,  266 
Balsa  minaceae,  396,  586 
Baptisia  bracteata  Ell.,  392 

leucantha  T.  &  G.,  393 
Barbula  ruralis  Hedw.,  40 

tortuosa  W.  and  M.,  40 
Bartramia  oederi  Schw.,  42 

pomiformis  Hedw.,  42 
Batrachium  S.  F.  Gray,  460 

aquatile  Wimm.,  461 

circinatinn  Spach.,  460 

divaricatum    (Schrank)    Wimm., 
360,  388,  460 

hederaceum  S.    F.  Gray,  462 

Lobbii  Howell,  462 

trichophyllum     (Chaix)     Bossch, 

360,  388,  461 
Bazzania   tnlobata    (L- )   S.    F.    Gray, 

193 
Beckmannia  erucaeformis  (L.)   Host, 

574 

Brgonia,  54 
Berberidaceae,  388,  582 
Berchemia  racemosa  Sieb.  &  Zucc.,  78, 

85,87,  116,  13  r,  132,  133 
Berula    erecta    ( Huds. )    Coville,   360, 

400 


Betula,  429,  430,  565 

glandulosa  Michx.,  578 

lenta  L.  368,  381 

lutea  Micbx.,  225,  366,  381 

nigra  L.,  358,  381 

papyrifera  Marsb.,  366,  367,    381, 
578 

pumila  L.,  382 
Betulaceae,  381,  578 
Biatora,  6,  221,  227,  236,  282,  672,  681 

akompsa  Tuck.,  702 

atrogrisea  (Delis)  Hepp.,  671,  702 

arthropurpurea     (Mass.)     Hepp., 

267,  671,  701 

atropurpurea  (Mass.)  Hepp.,  672 
coarctata  (Sm.,  Nyl.)  Tuck.,  230, 

266,  323 

var.  brajeriana  Schaer.,  17 
var.  brugeriana,  323 
decipiens   (Ehrh.)    Fr.,    281,   296, 

322,  668,  669,  700 
var.   dealbata  Auct.,  281,  296, 

322,  668,  700 
flavidolivens  Tuck.,  268 
flexuosa  Fr.,  281,  323 
fossarum  (Duf. )  Mont.,  296 
fuscorubella  (Hoffm.)  Tuck.,  230 

268,  291,  303,  304,  323,  671,  702 
glauconigrans    Tuck.,     228,    267 

680,  702 
hypnophila    (Turn.)    Tuck.,    268, 

323,  680,  702 
icterica  Mont.,  293,  322 
incompta  (Borr. )  Hepp.,  268 
inundata  Fr.,  298,  324,  666,  702 
leucophaea    (Floerk. ),    267,    662, 

700 

var.  griseoatra  Koerb.,  228,  267 
lucida  (Ach.)  Fr.,  228.  268 
mixta  Fr.,  701 

var.  atlantica  Tuck.,  701 
muscorum  (Sw. )  Tuck.,  230,  269, 

286,  293,  296,  324,  668,  669,  703 
myriocarpoidts  (Nyl.)  Tuck.,  17, 

230,  268,  289,  323,  701 
naegelii  Hepp.,  230,  268,  304,  323 
oxyspora  (Tul.)  Nyl.,  229,  268 
prasina  Fr.  var.  byssacea  Th.  Fr., 

701 
rubella    (Ehrh.)     Rahenh.,     268, 

303,  323.  67i,  672,  680,  702 


750 


INDEX. 


Biatora 

rufonigra  Tuck.,  3,    17,   266,   282, 

287,  288,  289,  322,  665 
russellii  Tuck.,  297 
sanguineoatra  (Fr. )  Tuck.,  267 
schweinitzii  Fr.,  229,  268 
sphseroides  (Dicks.)   Tuck.,   230, 

267,  680,  702 
subfusca  Fr.,  303 
suffusa  Fr.,  281,  323,  702 
trachona  Flot.,  299 
turgida  (Fr. )  Nyl.,  227 
turgidula  (Fr. )  Nyl.,  267,  671,  700 
uliginosa  (Schrad. )  Fr.,  267,  294, 

323,  673,  701 

varians  (Ach.)  Tuck.,  323,  680,  701 
vernalis  (L.)  Fr.,  266,  680,  700 
viridescens    (Schrad.)    Fr.,     266, 

679,  700 
Bicuculla  cucullaria  (L. )   Millsp. ,  362, 

388,  555 
Bidens  cernua  L,.,  597 

comosa   (A.  Gray)  Wiegand,  358, 

362,  368,  414 
frondosa   L.,    358,    362,   414,   425, 

597,  626 
laevis  (L.)  B.S.P.,   358,  4M,  597, 

626 

Bignoniaceae,  82,  124 
Blepharostoma     trichophyllum     (L. ) 

Dutnort,  193 

Blephila  hirsuta  (Pursh)  Torr.,  404 
Boltonia  asteroides  (L. )  L'Her.,  411, 

626 

Boraginaceas,  403,  590 
Botrychium  lunaria  (L-  ~)  Sw.,  362,  369, 

619 

virginianum  (L. )  Sw.,  616 
Botrydium,  198 
Botryococcus,  614 
Boutelouacurtipendula  (Michx.  )Torr., 

367,  375,  574 
hirsuta  Lag.,  367,  375 
oligostachya  (Nutt. )  Torr.,  21,  22, 

574 
Brachyactis  angustus  (Lindl. )  Britton, 

596 

Brachypuccinia,  543,  548 
Brachythecium,  36 

campestre  Sch.,  49 

flexicaule  Ren.  and  Card.,  49 


Brachythecium 

orycladon  (Brid.)  Grout,  49 
plumosum  Sch.  (?),  49 
salebrosum  Sch.,  49 
starkei  Sch.,  49 
Brassica  arvensis  (L. )  B.S.P.,  389 

nigra  (L.)  Koch,  388,  622,  627,  628 
Brauneria  pallid  a,  20,  21 
Bromus  ciliatus  L.,  375,  549,  574 
kalmii  A.  Gray,  375,  574 
purgans  L.,  574 
secalinus  L.,  375 
Broussonetiapapyrifera  (Linn.)  Vent., 

?*•  85,  87,  94,  130,  131,  132,  133 
Buellia,  6,  221,  227,  282,  283,  665,  672, 

674 
alboatra    (Hoffm.)    Th.    Fr.,   270, 

303,  324,  671,  704 
var.  saxicola  Fr.,  281,  298,  299, 

324 

dialyta  (Nyl.)  Tuck.,  230,  271 
geographica  (Pers. ). Tuck.,  232,  271 
myriocarpa    (DC.)     Mudd. ,    230, 

271,  324,  673,  704 
var.    polyspora   Willey,    230, 

271,  704 

parasema  (Ach.)  Th.  Fr.,  270,  303, 
306,  324,  671,  672,  673,  675, 
704 

var.  triphragmia  Nyl.,  271 
parmeliarum    (Sommerf. )   Tuck., 

230,  272 

petrsea  (Plot.,  Koerb.)  Tuck.,  3, 

17,  223,  224,  233,  271,  282, 

287,  289,  325,  662,  664,  704 

var.  grandisFloerk.,17,229,271 

var.   montagnaei,  17,  233,  271, 

285,  325,  664,  704 
pullata  Tuck.,  278,  281,  282,  285, 

287,  325,  664,  704 
spuria  (Schoer. )  Arn.,  17,  285,  287, 

289,  324 
turgescens  (Nyl.)  Tuck.,  281,  306, 

325 

Bursa  bursa-pastoris  (L.)  Britton,  137, 

142,  389 
Butneria  fertilis  (Walt.)  Kearney,  73, 

85 
florida  (Linn.)  Kearney,  72,  85,  87, 

loo,  130,  131,  132,  133 
Butomineae,  651 


INDEX. 


751 


Cactus  viviparus  Nutt.,  192 
Ccsotna  nitens  Schwein.,  552 

(^Ecidium )  pedatatum  Schw.,  632 
(sEcidium)  sagittatum  Schw.,  632 
Csesalpinaceae,  73,  102,  392 
Calamagrostis      langsdorfii       (Link.) 

Trin.,  620 

lapponica  Trin.,  620 
Caliciuui,  225,  236,  663,  673.  674,  678, 

679,  706 

brunneolum  Acli.,  273 
chrysocephalum      (Turn.)     Ach., 

273,  677,  678,  706 
var.  filare  Ach.,  229,  273 
curtum  Turn,  and  Borr.,  678,  706 
hyprcllum  Ach.  var.  viride  Nyl., 

274,  671,  673 
lucidum  (Th.  Fr.)  Fink.,  659,  661, 

675,  676,  706 
paiietinum    Ach.,    273,    306,    326, 

673.  679.  707 
polyporaeuni  Nyl.,  706 
pusillum  Flk.,  707 
quercinum  Pers.,  274,  306,  326,  694 
trabinellum   (Schaer. )   Kbr.,  673, 

679.  707 

trachelinum  Ach.,  678,  706 
trichiale  Ach.,  273 

var.   cinereum   Nyl.,  673,  677, 

678,  706 
var.  stemoneum  Nyl.,  230,  273, 

707 

turbinatum  Pers.,  274,  695 
Calla  palustris  L.,  576 
Calliblepharis,  156,  205 
Calothrix  parietiua  (Nag.)  Thur.,  27 
Caltha,  500 

palustris  L.,  36ri  387,  544 
Calycanthaceae,  72,  100 
Calycanthus  occidentalis,  101 
Calyptospora  J.  Kiihn,  537,  540 

goeppertiana  Kuhn,  540 
Camclina  sativa  (L. )  Crantz.,  389 
Campanulaceae,  408,  593 
Campanula  americana  L-,  362,  408 
aparinoides  Pursh,  408,  593 
rotundifolia  L.,  367,  408,  593 
Camptosorus  rhizophyllus  (L- )  Link 

367.  370 

Cannabis  sativa  L.,  383 
Capea,  723 


Capparidacese,  389 
Caprifoliaceae,  83,  407,  592 
Cardamine  bulbosa  (Schreb. )  B.S.P., 

360,  389 
Carduus,  425,  548 

altissimus  L.,  598 

arvensis  (L. )  Robs.,  598,  627 

discolor  (  Muhl.)  Nutt.,  415,  598 

lanceolatus  L.,  415,  627,  628 

odoratus  (Muhl.)  Porter,  415 

undulatus  Nutt.,  598 
Carex  abacta  Bailey,  620 

castauea  Wahl.,  547 

cristatella  Brittou,  377,  576 

filiformis  L.,  576 

fusca  All.,  576 

hystricina  Muhl.,  376 

limosa  L.,  620 

lupulina  Muhl.,  376 

lurida  Wahl.,  368,  376 

retrorsa  Schwein.,  576 

rosea  Schk.,  362,  377 

stipata  Muhl.,  376 

sychnocephala  Carey,  576 

torta  Boott,  368,  376 

tribuloides  Wahl.,  377 

utriculata  Boott,  547,  576 

vulpinoidea  Michx.,  377 
Carpinus  caroliniana  Walt.,  362,  381 
Caryophyllacese,  89,  386,  581 
Cassia  chamaecrista  L-,  392 
Castalia  rigida,  357 

tuberosa  (Paine)  Greene,  356,  386, 

626 

Castilleja  acuminata  (Pursh)  Spreng., 
620 

coccinea  (L. )  Spreag.,  366,  406 

sessili flora  Pursh,  406 
Catalpa  catalpa,  127 

speciosa  Warder,  82,  85,  87,   126, 

130,  131,  132,  133 
Caulophyllutn        thalictroides       (L. ) 

Michx.,  362,  388,  582 
Ceanothus  americanus  L.,  396,  430 

ovatus  Desf.,  396 
Celastraceae,  76,  395 
Celastrus  scandens  Linn.,  76,  395 
Celtis,  130 

australis,  92 

occidentalis  L.,  70,  85,  87,  91,  130, 
13',  T32.  133.  366,  383 


752 


INDEX. 


Cenchrus  tribuloides  L,.,  375 
Cephalauthus  occidentalis   Linn.,   83, 
84,  87,  128,  130,  131.  132,  133,  358,  407 
Cephalozia    eaten  ulata    (Hiiben.) 

Spruce,  193 
media  Lindb.,  193 
Cerasterias,  614 
Cerastium  longipedunculatum  Muhl., 

361,  362,  386 
Ceratium  lonyicorne  Carter,  613 

tripos,  617 

Ceratodon  purpureus  Brid.,  40 
Ceratophyllacese,  386,  581 
Ceratophyllum  demcrsumL.,  196,  197, 

355,  386.  581 
Ceratosanthus  ajacis  Schur.,  435 

consolida  Schur.,  435 
Cercis   canadensis   Linn.,   73,    85,  87, 

104,  130,  131,  132,  133 
siliquastrum  Willd.,  73,  105 
Cetraria,  236 

aurescens  Tuck.,  241 

ciliaris    (Ach.)    Tuck.,    241,    306, 

309  659,  675,  677,  678,  683 
islandica  (L.)  Ach.,  230,  241 
juniperina  (L. )  Ach.,  676 

var.  pinastri  Ach.,    242,    663( 

665,  671,  677,  683 
lacunosa  Ach.,  241 
saepincola  (Ehrh.)  Ach.,  242 
Chsetochloa  viridis  (I,-)  Scribn.,  573 
Chantransia  expansa  Wood,  25 

pygmaea  (Kg.)  Sirdt.,  25 
Chamserhodos  erecta  (L. )  Bunge,  569, 

584 
Chara  contraria  A.  Br.,  25 

foetida  A.  Br.,  25 
Characeae  Richard,  25 
Characium  A.  Br.,  196 
Cheilanthesgraci1is(Fe£)Mett.,  368,371 

lanosa  (Michx.)  Watt.,  371 
Cheilosporum    calif ornicum    (Dene.), 

700,  702,  703 
frondescens    Post,  et   Rupr.,  700, 

702,  705 

f.  intermedia  Yendo,  704 
f.  maxima  Yendo,  704 
f.  polymorpha  Yendo,  704 
f.  typica,  703 

MacMillanii  sp.  nov.,  Yendo,  700, 
702,  706 


Cheilosporum 

planiusculum  (Ku'tz.),  700,  70* 
Chelone  glabra  L.,  361,  405 
Chenopodiacese,  89,  385,  579 
Chenopodium,  627,  628 

album  L.,  30,  579 

ambrosioides  L.  ,  569,  580 

Boscianum  Moq.,  30 

botrys  L.,  385 

glaucum  L,.,  580 

hybridum  L.,  580 

leptophyllum    (Moq.)    Nutt.,  580 
Chlorochytrium,  195,  196,  200 

archerianum  Hieron.,  26,  199 

cohnii  Wright,  196,  201 

dermatocolax,  199,  203 

inclu^um  Kjellman,  186,  198,  [99 
202,  203 

knyanum,  197 

laetum,  199 

lemnse,  196,  197,  198,  199,  203 

pallidum  Klebs,  197 

rubrum,  199 

schmitzii,  203 

viride,  199 

Chlorocystis,  199,  203 
Chlorophycese,  610,  614 
Chroococcus,  28,  614 
Chrysanthemum    leucanthemum    L., 

414 
Chrysomyxa  linger.,  537,  538 

ledi,  559 

pirolatum  (Koern.)  Wint.,  538 
Chrysopogon      avenaceus      (Michx.) 

Benth.,  374,  573 
Chrysopsis hispida  (Hook.)  Nult.,  569, 

595 

Chytridese,  198 

Cichoriacese,  409,  593 

Cichorium  intybus  L.,  409 

Cicuta  bulbifera  L.,  358,  361,  400,  588 
maculata  L-,  361,  399,  627 

Circaea,  550 

alpina  I,.,  366,  398,  550 
lutetiana  L-,  362,  398,  550 

Cistacese,  397,  586 

Cladonia,  221,  225,  668,  675,  683 

amaurocrsea    (Fl.)    Schaer.,   233, 

265 

bacillaris  Nyl.,  674,  700 
botrytes  (Hog.)  Willd.,  696 


INDEX. 


753 


Cladonia 

caespiticia  (Pers. )  Fl.,  16,  229,  262, 

264    301,  320,  321 
cariosa  (Ach.)   Spreng.,    16,    262, 

295,  321,  667,  679,  697 
cenotea  (Ach.)  Schaer.,  679,  698 
cornucopioides  (L.)  Fr.?  265,  301 
cornuta  (L. )  Fr.,  264,  667 
cristatellaTuck.,  220,  266,  306,  322, 

674.  700 

var.  paludicola  Tuck.,  281,  322 
decorticata  Floerk.,  230,  262 
defortnis   (L. )    Hoffm.,    227,    265, 

662,  699 

dejjenerans  Flk.,  263,  667,  698 
delicata    (Ehrh.)     Fl.,     17,     230, 

284 
digitata    (L.)    Hoffm.,    229,    265, 

662,  700 
fimbriata   (L.)  Fr.,  228,  262,  286, 

294,  295,  296,  321 

var.  apolepta  (Ach.)  Wainio, 

698 
var.  ceratodes  (Flk.)  Wainio, 

698 

var.  fibula  Ach.,  697 
Tar.  radiata  Fr.,  263,  321 
var.    simplex    (Weis. )    Plot., 

674,  697 

var.  subulata  (L.)  Wainio,  697 
var.   tubaeformis  Fr.,   16,  262, 

286,  295,  306,  321,  674,  678, 

697 
furcata    (Huds.)     Fr. ,    220,    223, 

264,  301,  322,  667,  679,  699 
var.  crispata  Fl.,  264 
var.  paradoxa  Wainio,  699 
var.  pungens  Fr.,  322 
var.  racemosa  Fl.,  301,  322 
var.  racemosa  Fr.,  301 
var.  scabriuscula  (Del.)  Ceon., 

699 
gracilis(L-)  Fr.,  16,  222,  263,  294, 

295,  306,  321,  667,  674,  678, 
698 

Nyl.,  220 

var.  anthocephala  Flk.,  662, 

698 
var.   cervicornis  Floerk.,  228, 

263 
var.  elongata  Fr.,  667 


Cladonia 

gracilis   (L. )    Nyl.,    var.    hybrids 
Schaer.,  263,  321,  667,   674, 

678,  679,  698 

var.  syuiphycarpia  Tuck., 
227,  263,  321,  667,  674 

var.  vertidllata  Fr.,  17,  263, 
294,  295,  306,  321,  674,  677, 

679,  698 

macilenta     (Ehrh.)     Hoffin.,    17, 

266,  306,  322,  674,  700 
mitrula  Tuck.,   16,   230,  262,  295, 

320,  674,  678,  696 

pyxidata   (L. )    Fr.,    16,   222,  223, 
262,  286,   293,  294,  295,  321, 
667,  674,  679,  697 
var.  chlorophaea  Flk.,  697 
var.    neglecta    (Flk.)    Mass., 

674.  697 

rangiferina  (L,.)  Hoffm.,   17,  220, 

222, 223. 264, 322, 667,  679,699 

var.    alpestris    (I,.)     17,   222, 

265,  667,  699 
var.    sylvatica    L.,    222,    223, 

265,  322,  699 
squamosa  Hoffin.,  16,  264,  699 

var.  phyllocomaRabenh.,  227, 

264 

symphycarpa  Fr.,  262 
symphycarpia  Tuck,  295,  306 
symphycarpia   Fr.  var.    epiphylla 

(Ach.)  Nyl.,  228,  262,  320 
turgida  (Ehrh.)  Hoffm.,  263,  294, 

321 
var.  conspicua  (Schaer.)  Nyl., 

263 
uncialis   (L.)   Fr.,   222,    223,    265, 

667,  699 
Cladophora,  614 

lanosa,  195 
Clathrocystis,  614 
Claytonia,  639 
Clematis  virginiaua  L,.,  362,  387,  427, 

555,  58i 

Climacium  americanum  Brid.,  48 
Clinopodium  vulgare  L.,  404 
Clintonia  borealis  (Ait.)  Raf.,  551 
Closterium  parvulum  Naeg.,  612,  614 
Cnicus,  425 
Coelastrum    microporum     Naeg.    var. 

speciosum  Wolle,  611,  614 


754 


INDEX. 


Ccelosphae  Hum  kiitzingiai  uin  Naeg., 

611,  612,  614 
Coccochloris,  28 
Coleosporium  Leveille",  537,  538 

sonchi-arvensis  (P.)  Wmt.,  539 
Collema,  195,  221,  282,  665,  668,  672 

crispum  Borr.,  667,  690 

flaccidum  Ach.,   15,  229,  254,  292, 
303,  314,  671,  680,  689 

furvum  (Ach.)  Nyl.,  254,  290,  292, 

299.  315 

limosum  Ach.,  655,  690 
marescens  (Huds. )  Ach.,  680 
nigrescens  (Huds.)  Ach.,  254,671, 

689 

plicatile  Schaer.,  281,  298,  299,  314 
pulposutn  (Barnh. )  Nyl.,  15,  294, 

314,  667,  690 

pustulatum  Ach.,  281,  298,  299,  314 
pycnocarpum  Nyl.,  230,   254,  303, 

314,  671,  689 
ryssoleum  Tuck.,  690 
tenax  (Sw. )  Ach.,  281,  294,  314 
Comaudra  utnbellata  (L. )   Nutt.,   20, 

384.  558 

Comarum  palustre  L.,  583 
Commelinaceae,  377 
Compositae,  410,  594 
Coniferse,  89 
Coniocybe,  236,  660 

pallida  (Pers. )  Fr.,  231,  239,  274, 

303,  326,  659,  695 

Conringia  orieutalis  (L-)  Dumort.,  191 
Constantiuea,  180,  182,  201,  203 
reniformis,  175,  176,  177,  178 
rosa-marina,    175,     176,    177,    178, 

185,  1 86,  187 
schmitzii,  200 
sitchensis,  175.   176,  177,  178,  186, 

187,  200,  202 
thiebauti,  178,  187 
Convallariaceae,  378,  577 
Convolvulaceae,  402,  590 
Convolvulus,  89 

sepium  L.,  402,  544,  590 
spithamaaus  L. ,  402,  544 
Corallina  aculeata  Yendo,  700,  702,  708 
frondescens  Post,  et  Rupr.,  703 
officinalis  L.,  707 

var.  chilensis  Kiitz,  700,  702, 
706,  707 


Corallina 

officinalis  L.  f.  <5  Yendo,  707 

pilulifera,  702 

planiuscula  Kiitz,  705 

squamata,  702 

tuberculosa    Post,    et    Rupr.,  701, 
702 

vancouveriensis  Yendo,  700,   702, 

707 

f.  densa  Yendo,  707 
f.  typica  Yendo,  707,  708 
Coreopsis  palmata  Nutt.,  367,  413,  426 
Cornacese,  81,  400,  588 
Corn  us,  564 

amonum  Mill.,  81,  358,2400 

canadensis  L.,  551 

candidissima  Marsh,  362,  400,  588 

circinata  L'Her.,  400 

florida  Linn.,  82 

rotundifolia,  362 

stolonifera   Michx.,    8;,  362,   400, 

588 

Corylus  americana  Walt.,  362,  381,  430, 
578 

rostrata  Ait.,  381 
Cosmarium,  617 

nitidulum  De  Not.,  612 
Costaria  turneri  Grev.,  725 
Crassulaceae,  387,  582 
Crataegus,  239,  429 

coccinea  L. ,  391,  584 

macracautha  Lodd.,  368,  391 

punctata  Jacq.,  362,  368,  391 

tomentosa  L.,  362,  391 
Crocus,  55,  62,  63,  65 
Cronartiuin  Fries.,  537,  538 

asclepiadeum     (Willd.)    Fr.     var. 

quercuum  B.  &  C.,  538 
Cruciferse,  89,  388,  582 
Cryptogramme  acrostichoides  R.  Br., 

619 

Cucurbitaceae,  88,  593 
Cuscuta  coryli  Engelm.,  402 

gronovii  Willd.,  402,  590 

indecora  Choisy,  402 

paradoxa  Raf.,  402 

polygonorum  Engelm.,  590 
Cuscutacese,  402,  590 
Cyanophyceae,  610,  614 
Cyclotella  comta  (Ehr.)  Kg.,  613,  614 
Cyliudrothecium  seductrix  Sulliv.,  48 


INDEX. 


755 


Cytnbella  lanceolata  (Ehr. )  Kirchn., 

613,  614 

Cynodontium  polycarpum  B.  S.,  37 
Cyperacete,  375,  575 
Cyperus,  627 

diandrus  Torr.,  575 

esculentus  L-,  357,  375 

filiculmis  Vahl.,  367,  376 

houghtoni  Torr.,  367,  368,  376 

schweinitzii  Torr,  367,  375,  376 

speciosus  Vahl,  575 
Cypripedium  candidum  Willd.,  379 

hirsutum  Mill.,  362,  366,  379 

reginae  Walt.,  379 

spectabilis,  366 
Cyrtorhyncba  Nutt.,  501 

ranunculina  Nutt.,  502 

Cymbalaria  Britton,  503 
Cystopteris  bulbifera  (L.)  Bernh.,  362, 

367,  3?o 
fragilis  (L,.)  Bernh.,  366,  370,  560 

Daldinia,  564 

concentrica  (Bolt.)  C.  &  N.,  567 
tuberosa  (Scop.)  Voss.,  567 
vernicosa   (Schw.)    Cesati    &    de 

Notaris,  567 
Dasystoma       grandiflora        (Benth.) 

Wood,  368,  406 
Datura  tatula  L,.,  405 
Delphinium  aconitifolium   Muhlenb., 

441 

albescens  Rydb.,  446 
alpinum  Waldst.  &  Kit.,  447 
apiculatum  Greene,  440 
Andersonii  Gray,  441 
azureum  Michx.,  444,  445 
azureum  Torr.,  451 
azureum  Torr.  &  Gray,  439 
var.  laxiflorum  Huth,  444 
var.  simplex  Huth,  451 
Barbeyi  Huth,  449 
bicolor  Nutt.,  438 

var.  cognatum  n.  var.  Davis, 

438 

var.  glareosum  Davis,  439 
var.  Montanense  Rydb.,  438 
var.  Nelsonii  Davis,  438 

bicornutum  Hemsl.,  452 

var.  Hemsleyi  Huth,  453 

Blochmana  Greene,  440 


Delphinium 

Brunoniannm  Royle,  443 

Californicum  Torr.  &  Gray,  437 
var.  laxiusculum  Huth,  437 
var.  scapigerum  Huth,  437 

camporum  Greene,  445,  446 

var.  macroseratilis  Davis,  446 
var.  penardi  Davis,  446 

Carolinianum     Walt.    Fl.    Carol., 

445,  446 

var.  album  Hort.,  445 
var.  vimineum  Gray,  445 

Cashmirianum  Royle,  443 
var.  Walkeri  Hook.,  443 

cardinale  Hook.,  436 

cheilauthum  Fisch.,  448 

Clusianum  Host.,  447 

coccineum  Torr.,  436 

Columbianum  Greene,  450 

consolida  Linn.,  419,  435 

dazniricum  Georgi,  452 

decorum  Beuth.,  440 

decorum  Fisch.  &  Mey.,  439 
var.  gracilentum  Davis,  439 
var.   Nevadense    Wats.,   441, 

442 

var.  nudicaule  Huth,  436 
var.  patens  Gray,  439 

discolor  Fisch.,  447 

distichum  Geyer,  451 

diversifolium  Greene,  449 

var.  diversifolium  Davis,  449 

Ehrenbergi  Huth,  453 

elatum  Linn.,  447 

var.  occidentals  Wats. ,  449 

Emilia  Greene,  441 

exaltatum  Alton,  443,  447 

exaltatum  Hook.,  448 

exaltatum  Hook.  &  Arn.,  437 
var.  Barbeyi  Huth,  449 
var.  Californicum  Huth,  437 
v&r.glaucum  Huth,  449 
var.  Nuttallii  Huth,  450 
var.  scopulorum  Huth,  448 
var.  trolliifolium  Huth,  447 

fissum  Waldst.  &  Kit.,  452 

flammeum  Kellogg,  436 

flexuosum  Raf.,  441 

formosum  Boiss.  &  Huet,  451 
formosum  Hort.,  448 

geraniifolium  Rydb.,  446 


756 


INDEX. 


Delphinium 

Geyeri  Greene,  445 

var.  geraniifolium  Davis,  446 
var.  Wootoni  Davis,  445 

glareosum  Greene,  439 

glaucescens  Rydb.,  450 

var.  multicaule  Rydb.,  450 

glaucum  Wats.,  449 

gracilentum  Greene,  439 

grandiflorum  Linn.,  448 

var.  Chinensis  Fischer,  448 
var.     variegatum     Hook.    & 
Arn.,  440 

Hanseni  Greene,  440 

Hanseni    var.  arcuatum   Greene, 
440 

hesperium  Gray,  439 

hespetium  Huth,  439 

var.  Hanseni  Greene,  440 
var.  recurvatum  Davis,  440 

hirsutum  Pers.,  452 

hybrid  urn  Steph.,  452 

var.  sulphureum  Hort.,  437 

intermedium  Willd.,  447 

latisepalum  Hemsl.,  453 

leptophyllum  Hemsl.  454 

leucophceum  Greene,  450 

lilacinum  Willd.,  443 

Maackianum  Regel,  451 

macroseratilis  Rydb.,  446 

Madrense  Wats.,  452 

magnificum  Paxt.,  448 

Menziesii  DC.,  442 

Menziesii  Gray,  438 

Menziesii  Wats.,  441 

var.  ochroleucum  Torr.  &  Gray, 

439 

var.  pauciflorum  Huth,  442 
var.  Utahense  Wats.,  438 

monophyllum  Gilib.,  435 

moschatum  Munro,  443 

Nelsoni  Greene,  438 

nudicaule  Torr.  &  Gray,  418,  436 

Nuttallianum  Pritz.,  442 

Nuttaliii  Gray,  450 

var.  leucophaeum  Davis,  450 

occidentale  Wats.,  417 

occidenta/eWats.,  449 

Oreganum  Howell,  445 

ornatum  Greene,  440 

palmattfiduin  DC.,  447 


Delphinium 

Patens  Benth.,  439 

pauciflorum  Nutt.,  442 

var.  depauperatum  Gray,  442 
var.  Nevadense  Gray,  442 

Parishii  Gray,  441 

Parryi  Gray,  441 

pauperculum  Greene,  442 

pedatisectum  Hemsl  ,  453 

peltatum  Hook.,  436 

Penardi  Huth,  446 

Przewalskianum  Hort.,  437 

Przewalskii  Huth,  437 

pubescens  Griseb.,  435 

pyramidale  Royle,  447 

ranunculifolium  Wall.,  447 

recurvatum  Greene,  440 

sarcophyllum  Hook.  &  Arn.,  436 

scaposum  Greene,  446 

scopulorum  Gray,  448,  449 

scopulorum  Wats.,  449 

var.  attenuatum  Jones,  449 
var.  glaucum  Gray,  449 
var.  stachydeum  Gray,  449 
var.  subalpinum  Gray,  449 

segetum  Lam.,  435 

simplex  Dougl.,  451 

simplex  Nutt.,  450 

simplex  Wats.,  439 

var.  distichiflorum  Hook.,  451 

sinense  Fisch.,  448 

sonnei  Greene,  441 

speciosum  Boiss.  &  Huet.,  451 

staphisagria,  417 

tauricum  Pallas,  452 

tenuisectum  Greene,  453 

Treleasei  B.  F.  Bush,  446 

tricorne  Michx.,  441 

var.  Andersonii  Huth,  441 
var.      depauperatum      Huth, 

442 
var.  patens  Huth,  439 

trollifolium  Gray,  419,  447 

trydactylum  Michx.,  443 

uliginosum  Curran,  447 

urceolatum  Jacq.,  454 

variegatum  Torr.  &  Gray,  440 
var.  apiculatum  Greene,  440 
var.  Blochmanae  Davis,  440 
var.  Emiliae  Davis,  441 

versicolor  Salisb.,  435 


INDEX. 


757 


Delphinium 

vimineuHi  D.  Don,  445 

virescens  Gray,  445 

virescens  Nutt.,  446 

virescens  Rydb.,  437 

virgatum  Jacq.,  448 

viride  Wats.,  438 

viridescens  Leiberg,  436 

Wislizeni  Engelni.,  454 

Wootoni  Rydb.,  445 

Zalil  Ait.  &  Hems.,  437 
Deringa  canadensis  (I,.)  Kuntze,  362, 

400,  588 

Descurainia  pinnata  Britton,  30 
Dtanthus  caryophyllus  L,.,  543 
Diatomaceae,  610 
Diatomese,  614 
Diueoma  Viola:  Kuntze,  633 
Dichelyma  pallescens  B.  S.,  44 
Dichothrix  calcarea  Tilden,  27 
Dicrauuin,  36 

boryeani  De  Not.,  37 

drummondii  C.  Mull.,  37 

flagellare  Hedw.,  38 

fuscescens  Turn.,  38 

longifolium  Hedw.,  38 

montauum  Hedw.,  39 

palustre  La  Pyl,  37 
alatum  Barnes,  37 

scoparium  Hedw.,  39 

undulatum  Ebrb.,  39 

viride  B.  S.,  39 
Dictyophora  duplicata,  534 

Ravenelii  Burt.,  525,  533,  534 
Dictyosphaerium     pulchellum    Wood, 

612,  617 
Diervilla   diervilla    (L.)   MacM.,   362, 

408 

Dilsea,  202 
Ditnorphococcus,  617 
Dioscorea  villosa  L,.,  379 
Dioscoreaceae,  379 
Dirca  palustris  L,-,  366,  398,  558 
Distichiutn  capillaceum  B.  S.,  40 
Dodecatheou  meadia  L.,  400 
Doellingeria  umbellata    (Dill.)   Nees, 

361,  539 

umbellata  pubens  (A.  Gray)  Brit- 
ton,  412,  596 

Dondia  depressa  (Pursh)  Briiton,  580 
Draba  caroliuiana  Walt.,  367,  389 


Dracocephalum  parvifloruui  Nutt.,  591 

Drupaceae,  584 

Drymocallis  arguta  (Pnrsh)  Rydberg, 
583 

Dryopteris  fragrans  (L- )  Schott,  619 
goldieana  ( Hook. )  A.  Gray,  368, 370 
spinulosa  (Retz.)  Kuntze,  370 
thelypteris  (L.)  A.  Gray,  361,  370 

Echinopanax  horridum  (Smith)  Dec. 

&  Planch.,  620 
Ecklonia,  723,  730 
Elaeagnaceae,  80,  121,  587 
Elaeagnus,  122 

argentea  Pursh,  587 

umbellata  Tbunb.,  80,  87,  121,  130, 

131,  133 
Eleocharis  acicularis    (L. )    R.   &  S., 

356,  357,  376,  575 
intermedia  (Muhl. )  Schultes,  575 
Elodea,  54 
Elymus  canadensis  L-,  367,  375,  575, 

627,  628 

macouni  Vasey,  569,  575 
virgiuicus  L.,  357,  375,  575 
Encalypta  ciliata  Hedw.,  41 
Encyouema,  614 
Endocarpon,  221,  669 

arboreum  Schwein.,  281,  327 
fluviatile  DC.,  222,   223,  224,  274, 

327,  707 
hepaticum  Ach.,  18,  286,  293,  294, 

296,  327.  707 

miniaturh  ( L. )  Schaer.,  18,  227, 274, 

285,  290,  298,  299,    300,  326 

var.  complicatum  Schaer.,  274, 

285,  287,  289,  326 
pusillum    Hedw.,    297,    300,    327, 

670,  707 
var.  garovaglii  Kph.,  286,  293, 

294,  301,  327 

Endophyllacese,  537 

Endosphaera,  198 

Erysiphe  communis  (Wallr. )  Fr.,  426 

Ephebe,  6,  282 

pubescens  Fr.,  224,  253,  313 
solida  Born.,  3,  14,  222,  224,  253 

Epilobium,  360 

adenocaulon  Haussk.,  362,  398 
coloratum   Muhl.,    362,   366,   398, 
539.  587 


758 


INDEX. 


Epilobium 

lineare  Muhl.,  539,  587 

palustre,  627 
Equisetaceae,  371 
Equisetum  arvense  L.,  362,  371,  622 

hyemale  L.,  372 

laevigatum  A.  Br.,  372 

prateuse  Ehrh.,  372 
Eragrostis  hypnoides  (Lam.)    B.S.P. 

357,  374,  375 
Erechtites  hieracifolia  (I/.)  Raf.,  366, 

414,  425 

Ericaceae,  400,  588 
Erigeron  acris  droebachianus  (O.  F. 

Mueller)  Blytt.,  620 
annuus  (L.)  Pers.,  412,  556 
philadelphicus  L.,  596 
pulchellus  Michx.,  362,  412 
ramosus   (Walt.)   B.S.P.,   21,  412, 

596 
Eriocarpum       spinulosum       (Nutt. ) 

Greene,  22,  550 
Erysimum  cheiranthoides  L-,  389,  582 

pinnatum  Walt.,  30 
Erysiphe,  424 

aggregata  (Peck)  Farlow,  427 

cichoracearum  DC.,  425 

comtnunis,  427 

galeopsidis  DC.,  428 

graminis  DC.,  428 

Erythronium  albidum  Nutt,  362,  378 
Eudorina  elegans  Ehrenb.,  612,  617 
Eucalyptus,  85 

citriodora  Hook.,  81,  85 

corymbosa  Sm.,  81,  85 

globulus  Labill.,    80,  85,  87,    122 

131,  132,  133 
uhymenia,  176 
Kuonymus,  89 

atropurpureus  Jacq.,  362,  395 
Eupatorium  ageratoides  L,.,  362,  366, 
410,  426,  557 

altissimum  L.,  410 

maculatum  L. ,  594 

perfoliatum  L.,  410,  557,  595,  627 

purpureum  L.,  357,  410,  557 
Euphorbia,  540,  542 

corollata  L.,  395,  559 

cyparissias  L.,  395 

glyptosperma  Engelm.,  395,  542, 
586 


Euphorbia 

heterophylla  L.,  367,  395,  542 

maculata  L.,  395,  542,  586 

marginata  Pursh,  542 

nutans  Lag.,  395 

serpyllifolia  Pers.,  542,  586 
Euphorbiaceae,  395,  586 
Euphrasia  americana  Wettst.,  620 
Eurynchium   robustum  (Roell.)  Hol- 
zinger,  49 

strigosutn  Sch.,  50 
Euthamia  graminifolia  (L,. )  Nutt,  411, 

557,  595 

Euuromyces,  540 

Evernia  furfuracea  (L. )  Mann.,  242 
prunastri  (L.)  Ach.,  242,675,  677, 
684 

Fagaceae,  382,  578 

Falcata  comosa  (L. )  Kuntze,  362,  366, 

394,  427,  554,  585 
pitched   (T.  &  G.)  Kuntze,  368, 

394 
Ficaria  Huds.,  500 

ambigua  Bor.,  501 

aperata  Schur.,  501 

calth&folia  Reichb.,  501 

comtnunis  Dum.,  501 

Ficaria  Karst,  501 

grandiflora  Robert,  501 

Holubyi  Schur.,  501 

intermedia  Schur.,  501 

nudicaulis  Kern.,  501 

polypetala  Gilid.,  501 

ranunculoides  Moench.,  501 

Roberti  F.  Schultz,  501 

rotundifolia  Schur.,  501 

transsilvanica  Schur.,  501 

verna  Huds.,  501 
Fissidens  incurvus  Schw.,  40 

osmundoides  Hedw.,  40 
Fontinalaceae,  36 
Fontinalis  antipyretica  Linn.,  43 

duriaei  Sch.,  43 

holzingeri  Cardot,  43 

hypnoides  Hartm.,  44 

missourica  Card.,  43 
Fragaria   americana  (Porter)   Britton, 

362,  390 
Fragilaria,  611,  614 

capucina  Desmaz.,  613 


INDEX. 


759 


Fraxinus,  239,  274 

Americana  L.,  192,  559 
lanceolata  Borck.,  192,  358,  401 
nigra  Marsh.,  358,  401 

Freesin,  55,  62,  63 

Frcelichia  floridana  (Nutt. )  Moq.,  385 

Frullania  Eboracensis  Gottsche,  193 

Fucus  rosa-marina,  175,  187 

Funaria  hygromctrica  Hedw.,  42,  626 

Gaillardia  aristata  Pursh,  598 
Gileopsis  tctrahit  L.,  591 
Galtutn  apartne  L,.,  362,  407 

asprcllum  Michx.,  407,  544 

boreale  L.,  362,  407,  592 

concinnum  Torr.  &  Gray,  544 

trifidum  L.,  362,  407,  592 

triflorutn  Michx.,  362,  407 
Gaura  biennis  L  ,  368,  398 

coccinea  Pursh,  587 
Gentiana  acuta  Michx.,  589 

andrewsii  Griseb.,  544,  589 

crinita  Froel.,.36r,  401 

detonsa  Rottb.,  589 

flavida  A.  Gray,  361,  401,  589 

puberula  Michx.,  544,  589 

quinquefolia  L.,  401 

rubricaulis  Schwein,  620 
Gentianaceae,  401,  589 
Geraniaceae,  394,  585 
Geranium,  54 

bicknellii  Britton,  31,  585 

Carolinianum,  31 

maculatum  L.,  362,  394,  556 
Gerardia  aspera  Dough,  406,  592 

purpurea,  21 

tenuifolia  Vahl,  406,  592 
Geum  canadense  Jacq.,  391 

strictum  Ait.,  362,  391 

virginianum  L.,  584 
Gigartina,  154,  161,  165,  199,  205 

exasperata  Harv.,  601 
Glecoma  hederacea  L-,  404 
Gleditsia   triacanthos   L.,    73,   85,    87, 
105,  130,  131,   132,  133,  358,  368,  392 
Gloeocapsa  calcarea  Tilden,  29 
Gloeocystis,  617 

gigas  (Kg.)  Lagerh.,  612 
Gloeotrichia,  611,  614 

pisum  (Ag. )  Thuret.,  27 
Glycyrrhiza  lepidota  Pursh,  23,  585 


Gnaphalium  obtusifolium  L.,  412 
Gracilaria,  205 
Graminese,  374,  572 
Graphis,  225,  227,  660 
dendritica,  272 
scripta  (L.)   Ach.,   272,  303,   304, 

325,  671,  672,  705 
var.  limitata  Ach.,    229,    272, 

303,  325,  671,  705 
var.  recta  (Humh. )  Nyl.,  225, 

272,  325,  671,  705 
Grimmia  apocarpa  Hedw.,  40 
Grindelia   squarrosa    (Pursh)   Dunal., 

22,  192,  426,  595 
Grossulariaceae,  390 
Gunnera  scabra,  195 
Gyalecta,  236 

fagirola  (Hepp. )  Tuck.,  230,  261 
Gymnocladus  dioica  Koch,  366,  392 
Gyninoconia  Lagerh.,  537,  551 

interstitialis   (Schlect. )    Lagerh., 

552 
Gymnosporangium  De  Candolle,  537, 

553 

clavariaeforme   (Jacq.)   Rees.,  553 
globosum  Farl.,  553 
juniperi-virgianianse  Schw.,  553 
macropus  Lk, ,  553 
nidus  avisThaxter.,  553 

Habenaria  bracteata  (Willd.)  R.  Br., 

362,  379 
leucophsea  (Nutt.)  A.   Gray,  361, 

379 

psycodes  (L,.)  A.  Gray,  361,  379 
Haloragidacese,  587 
Halymenia  reniformis,  177 
Hamamelidaceae,  390 
Hamamelis  virginiana  L.,  368,  390 
Hedeoma  hispida  Pursh,  404 

pulegioides  (L.)  Pers.,  404 
Hedwigia  ciliata  Ehrh.,  40 
Heleniutn  autumnale  L-,  358,  414 

autumn  ale   pubescens  (Ait.)  Brit- 
ton,  597 

Heliantbemum  canadense  (L  )  Michx., 
397 

majus  (L.)  B.S.P.,  397 
Helianthus,  426,  543,  545,  546 

annuus  L-,  191,  546,  597 

atrorubens  L.,  368,  413 


760 


INDEX. 


Helianthus 

decapetalus  L.,  426 

divaricatus  L,.,  4  [3,  426,  557 

giganteus  L.,  546,  627 

grosse-serratus  Martens,  413,  426, 
546,  597 

maximiliani  Schrad.,  597 

occidentalis  Riddell,  367,  413 

rigidus,  20,  21,  22 

scaherrimus  Ell.,  413,  426,  597 

strumosus  L.,  413 

tuberosus  L.,  413,  426,  546,  597 

tracheliifolius  Mill.,  413 
Heliopsis   helianthoides   (It.)    B.S.P., 
412,  546 

scabra  Dunal,  412,  425,  596 
Helleborus,  500 
Heltninthocladiaceae  (Harv. )  Schmitz, 

25 

Helobiae,  651 
Hemipuccinia,  543,  549 
Hemiuromyces,  540,  543 
Hepatica  acuta   (Pursh)  Britton,  362, 

387 
Heppia,  657 

despreauxii  (Mont.)  Tuckm.,  281, 

293.  295,  296,  313,  656,  677 
polyspora  Tuck.,  281,  282,  295,  313 
Heracleurh  lanatum  Michx.,  362,  399, 

588 

Heter-eupuccinia,  543,  546 
Heterothecium,  236,  662 

sanguinariutn  (L. )  Plot.,  233,  269 

var.  affineTuck.,  228,  269 
Heuchera  hispida  Pursh,  389 
Hicoria  minima  (Marsh.)  Britton,  380 

ovata  (Mill.)  Britton,  366,  380 
Hieracium  canadeuse  Michx.,  367,  409, 

548,  556,  594 

scabrum  Michx.,  366,  409 
utnbellatum   L.,   366,  368,  409 
Hippuris  vulgaris  I/.,  587 
IfomaliajamesuSchimp.,  45,  46,  47 
macounii,  46,  47 
trichomanoides,  45,  46,  47 

jamesii  (Schimp.)  Holzinger, 

44,  47 
Homalocenchrus    virginicus   (Willd.) 

Britton,  357,  375 
Hordeuni  jubatum  L.  575,  627 
vulgare  L-,  548 


Hormiscia  zonata  (Web.  and  Mohr) 
Aresch.  var.  valida  (Nag.)  Rabenh., 
26 

Houstonia  longifolia  Gaertn.,  592 

Humulus  lupulus  L.  362,  383,  424,  578 

Hydrocotyle  americana  L.,  368,  400 

Hydrocytium,  196 

Hydrophyllaceae,  402 

Hydrophylluui    appendiculatum 

Michx.,  402 
virginicum  L,.,  362,  402,  426,  551, 

559 
Hylocomium  splendens  Sch.,  52 

triquetrum  Sch.,  52 
Hypericaceae,  397,  586 
Hypericum  ascyron  L.,  397 

maculatum  Walt.,  366,  397 

majus  (A.  Gray)  Britton,  366,  397 
Hypnaceae,  36 
H>pnutn  chrysophyllum  Brid.,  50 

crista-castrensis  L..  51 

cupressiforme  ericetorum  B.  S.,  50 

filicinum  aciculjnum   C.    M.   and 

K.,  51 
trichodes  Brid.,  51 

haldanianum  Grev.,  51 

hispidulum  Brid.,  51 

reptile  Rich.,  51 

schreberi  Willd.,  51 

uncinatum  Hedw.,  52 
Hypoxis  hirsuta  (L. )  Coville,  379 
Hypoxylon,  564 

annulatum  (  Schw. )  Montague,  566 

atropurpureum  (Fries)  Fries,  566 

commuiatum  Holwayanum  Sacc., 

565 

comtnutatum  Nitschke,  565 
epiphaeum  Berkeley  &  Curtis,  564, 

566 

epiphlceum  B.  &  C.,  566 
ferrugineum  Holway  &  Ellis,  566 
fuscum  (Pers. )  Fries,  565 
granulosum  Bulliard,  565 
marginatum  Berkeley,  566 
rnorsci  Berkeley  &  Curtis,  565 
multiforme  Fr.,  565 
perforatum  (Schw.)  Fries,  566 
petersii  Berkeley  &  Curtis,  565 
rubiginosum  Ellis  &  Everhart,  566 
rubiginosum    (Pers.)    Fries,    563, 

564,  566 


INDEX. 


761 


Hypoxylon 

serpens  (Pers.)  Pries,  567 

transversum  Scbw.,  565 
Hystrix  hystrix  (L. )  Millsp.,  575 

Ilysanthes    gratioloides   (L-)    Benth., 

358,  406 
Impatiens,  556 

aurea  Muhl.,  362,  396 

biflora  Walt.,  362,  549,  556,  586 
Iridacese,  379,  577 
Iris  versicolor  L-,  379,  554.  577 
Ithyphallus,  534 

impudicus,  534 

tenuis,  533,  534 
Iva  xanthifolia  (Fresen.)  Nutt.,  594 

Jania,  701 

Juglandaceae,  380 

Juglans  cinerea  L.,  362,  380 

nigra  L.,  362,  380 
Juncaceae,  377,  576 
Juncus  acuminatus  Michx.,  577 

articulatus  L,.,  620 

balticus  Willd.,  576 

dudleyi  Wicgand,  569,  576 

effusus  L.,  377,  626 

nodosus  L.,  577 

tenuis  Willd.,  377,  577 

torreyi  Coville,  577 

vaseyi  Engelm.,  576 
Jungertnannia  barbata  Schreb.,  193 

quinquedentata  Web.,  193 

vetitricosa  Dicks.,  193 
Juniperus     cotnmunis     I,.,    367,    372, 

553 

sabina  L.,  367,  368,  372 
virginiana  L.,  366,  372,  553 

Kallymenia,  205 

reniformis,  177 
Koeleria  cristata   (L. )   Pers.,   21,   367, 

375,  574 
Koellia     flexuosa      (Walt.)     MacM., 

591 

virginiana  (L. )  MacM.,  405,  545 
Kuhnia  eupatorioides  L.,  410,  549 

glutinosa,  20,  21,  22 
Kuhnistera  Candida  (Willd.)  Kuntze, 

20,  21,  23,  367,  393,  585 
occidentalis,  22 


Kuhnistera 

purpurea   (Vent.)  MacM.,  20,  21, 

367,  397,  585. 
Kumlienia  Greene,  500 

Cooleyce  Greene,  502 

histricula  Greene,  500 

Labiatae,  403,  590 
Laciniaria,  539,  557 

apycnostachya  (Michx.)    Kuntze, 

595 
cylindracea  (Michx.)  Kuntze,  367, 

410 

punctata  (Hook.),  Kuntze  595 
pycuostachya    ( Michx. )    Kuntze, 

410 
scariosa    (L. )   Hill,    21,    367,   410, 

595 

Lactuca,  546,  626,  628 
canadensis  L.,  557 
floridana  (L.)  Gaertn.,  362,  409 
ludoviciana  (Nutt.)  DC.,  368,  409, 

557,  593 

pulchella  (Pursh)  DC.,  546,  594 

sagittifolia  Ell.,  368,  409 

scaiiola  L.,  192,  409,  626 
Lagedia  oxyspora  (Nyl.)  Tuck.,  696 
Laminaria,  728,  730,  732 

japonica,  731 

radicosa,  731 

saccharina,  728 
Lappula,  426 

americana    (A.    Gray)    Rydberg, 

569,  59° 

floribuuda  (Lehm. )  Greene,  590 
lappula   (L.)  Karst.,  367,  403,  590 
virginiana  (I/.)  Greene,  403,  426 
Lathyrus,  427,  429,  430,  540 

ochroleucus  Hook.,  362,  393 
venosus  Muhl.,  362,  393,  427,  541 
Lecanora,  221,  236,  280,  282,  316,  665, 

670,  674 

atra  (Huds. )  Ach.,  15 
bookii  (Fr.)  Th.  Fr.,  281,  282,  319 
calcarea  (L,. )  Soinmerf.,  229,  259, 

297,  3i8 

var.  contorta  Fr.,  227,  259,  287, 
288,  297,  318,  664,  669,  694 
cervina  (Pers.)  Nyl.,  319,  664,  694 
var.   cinereo-alba    Fink,    282, 
287,  288,  319 


762 


INDEX. 


I/ecanora 

cinerea  (L,. )  Somtnerf.,  16,  223, 
224,  258,  284,  287,  289,  318, 
664,  693 

var.  cinereo-alba  Fink,  280 
var.  gibbosa  Nyl.,  258,  318 
var.  Isevata  Fr.,  16,  258,  318 
elatina  Ach.,  229,  258 
erysibe  Nyl.,  300,  318 
frustulosa  (Dicks.)  Mass.,  232,  256, 

278,  287,  288,  317 
fuscata  (Schrad.)  Th.  Fr.,  16,  259, 

287,  319,  664,  694 

var.  rufescens  Th.  Fr.,  259 
gelida  (L. )  Ach.,  316 
gibbosa   (Ach.)   Nyl.   var.   micro- 

spora  A.  Zahl,  694 
hageni  Ach.,  15,  228,  257,  287,  300)  , 

305,  318,  664,  692 
var.    sambuci    (Pers. )  Tuck., 

257 
muralis    (Schreb. )    Schaer.,    287, 

317,  692 

var.  diffracta  Fr.,  228,  256 
var.    saxicola     Schaer.,     256, 

285,  287,  317,  664,  692 
var.   vcrsi color  Fr.,    297,  317, 

657,  692 
pallescens  (L. )  Schaer.,  258,  678 

693 

pallida  (Schreb.)  Schaer.,  228,  256, 

678,  692 
privigna    (Ach.)    Nyl.,    297,    319, 

669,  670,  694 
var.  pruinosa  Auct.,  297,  319, 

669,  700 

rubina(  Vill. )  Ach.,  15, 223, 224, 256, 

284,  287,  289,  317,  652,  664 

var.  heteromorpha   Ach.,  256, 

284,  287,  289,  317 
sordida  (Pers.)  Th.  Fr.,  232,  257 
subfusca  (L,- )  Ach.,  15,  223,   257, 
258,  289,  303,  304,  317,  657, 
671,  660,  676,  692 
var.  allophana  Ach.,  280,  287, 

288,  317 

var.  argentata  Ach.,  317,  675, 

692 
var.  coilocarpa  Ach.,  15,   257, 

287,  288,  301,  318,  692 
var.  distans  Ach.,  318 


L,ecanora 

subfusca  (L,.)  Ach.,   var.    hypno- 

rum  Schaer.,  229,  257 
tartarea  (L.)  Ach.,  230,  258 
varia   (Ehrh.)   Nyl.    15,   257,   258, 
289,  305,  318,  660,   664,  671, 

672,  673,  675,  693 

var.  polytropa  Nyl.,  661,  664,. 

693 

var.  saepin cola  Fr.,  257,  673,693 
var.  symmicta  Ach.,   16,  257,. 

673,  693 

variolascens  Nyl.,  692 
verrucosa  (Ach.)  Laur.,  671 
xanthophana  Nyl.,   278,   284,  287,. 

289,  319,  664,  694 
Lechea  Leggettii,  32 
minor,  32 

stricta  Leggett,  32,  397,  586 
I^ecidea,  6,  221,  227,  236,  674,  703 
acclinis  Flot.,  229,  270,  703 
albocaerulescens  (Wulf. )  Schaer.,. 

3,  17,  229,  270 
crustulata  Ach.,  227,  269 
cyrtidia  Tuck.,  230,  270 
enteroleuca  Fr.,  270,  303,  304,  306,. 

324,  671,  672,  703 
var.    achrista  Sommerf.,   227,. 

270,  305,  324,  703 
var.  ambigua  Naz. ,  673,  703 
var.  flavida  Fr.,  703 
lactea  Fl.,  232,  269 
lapicida  Fr.,  229,  269 

var.  oxydata  Fr.,  228,  269 
melancheima  Tuck.,  270 
platycarpa  Ach.,  229,  270 
speirea  Nyl.,  270 
spirea  Ach.,  227 

Legouzia  perfoliata  (L.)  Britton,  408 
Leguminosae,  552 
lyejeuuea  serpyllifolia    (Dicks.)  Lib.>, 

193 
L,emna,  195 

minor  L.,  197,  355,  377,  57$ 

trisulca  L.,  576 
Lemnacese,  377,  576 
Lentibulariacese,  406 
Leonurus  cardiaca  L.,  404 
Lepachys  columnaris,  22 
Lepidium,  627 

apetalum  Willd.,  388 


INDEX. 


763 


Lepidozia  reptans  (L. )  Pumort,  193 
Leptandra   virginica    (L.)    Nutt.,  406, 

592 
Leptilon  canadeuse   (L. )  Britton,  412, 

596,  626 

Leptobryum  pyriforme  Scb.,  42 
Leptogium,  282,  660 

chloromelum   (Sw.)  Nyl.,  15,  299, 

30!.  315 
lacerutn   (Sw. )   Fr.,  223,  224,   230, 

254,  290,  2yS,  299,  315,  780 
var.    pulvinatutn   Moug.     and 

Nestl,  230,  254 

inyochroum     (Ehrh.,    Schaer. ) 
Tuck.,    229,    254,    290,   303, 
315,  671,  680,  690 
var.      tomeutosum      Schaer., 

255 

pulchellutn  (Ach.)  Nyl.,  280,  315 

tremelloides  (L. )  Fr.,  15,  254 
Leptopucciniu,  543,  550 
Leptorchis  liliifolia  (L,. )  Kuutze,  363, 

380 
Leskea,  36 

nervosa  Myr.,  47 

polycarpa  paludosa  Sch.,  47 
Lespedeza  capitata   Michx.,   393,  430, 

543 

violacea  (L. )  Pers.,  430 
Lesquerella  argcntca,  22 
Lessonia,  726,  728,  730,  732,  739 
Leucobrj  um  glaucum  Sch.,  40 
Liliacese,  378,  577 
Lilium  canadensc  L,.,  361,  378,  554 

philadclphicum  I,.,  378 

umbell  itum  Pursh,  366,  378 
Linaceae,  394,  585 
lyinum  rigidum,  21 

sulcatuin  Riddel  1,  367,  394,  585 
Lippia  lanceolata  Michx.,  358,  403 
Liriodendron   tulipifera   Linn.,  72,  85, 

87,  96,  130,  131,  132,  133 
Lithospermum  augustifolium  Michx., 

403 

canescens  (Michx.)  Lehm.,  403 
gmelini  (Michx.)  A.  S.  Hitchcock, 

403 

Lobelia  cardinalis  L.,  358,  408 
cordi folia,  366 
inflata  L.,  366,  408 
kalinii  L.,  593 


Lobelia 

spicata  Lam.,  367,  408,  593 
syphilitica  L.,  361,  363,  408,  593 

Lonicera,  430 

dioica  L.,  363,  408,  430 
hirsuta  Eaton,  430 
sullivantii  A.  Gray,  363,  408 

Loranthus  neelgherrensis  L.,  170,  172 

Lotus  americanus  (Nutt.)   Bisch.,  427, 
584 

Lupinus  perennis  L.,  554 

Lychnis  flos-cuculi,  199 

Lycodonum  sylvaticum  Fourr.,  349 

Lycopodium  selago,  L.,  619 

Lycopsis  arvensis  L.,  403 

americanus  Muhl.,   358,  405,  591, 

623,  625 

lucidus  Turcz.,  358,  405,  591 
rubellus  Moench,  358,  405 
virginicus  L.,  358,  405,  546,  591 

Lygodesmia  juncea,  22 

Lyngbya,  614 

majuscula  Harv.,  612 
martensiana  Menegh.var.  calcarea 

Tilden,  28 
nana  Tilden,  28 

Lyngbyese  Gomont,  28 

Lysimachiaterrestris  (L. )  B.  S.  P.,  400 

Lythraceae,  398 

Ly thrum  alatum  Pursh,  358,  361,  398 

Macrocalyx  nyctalea  (L- )  Kuntze,  361, 

403 

Macrocystis,  726 
Magnoliaceae,  72,  96 
Malus  coronaria,  31 

ioensis  .  Wood)  Britton,  31, 363, 391 
Malva  rotundifolia  L.,  396 
Malvaceae,  396 
Marchantia  polymorpha,  626 
Meibomia,  543 

canadensis  (  L.  )  Kuntze,  393,  430, 

585 

dilleuii  (Darl.)  Kuntze,  368,  393 
grandiflora  (Walt. )  Kuntze,  393 
illinoensis  (A.  Gray)  Kuntze,  368, 

393 

Melampsora  Castagne,  537,  539 
epilobii  (P.I  Fckl.,  539 
populina  (Jacq.)  Lev.,  539 
salicis-caprese  (P.)  Wint,  540 


764 


INDEX. 


Melampsoracese,  537,  538 
Melanthaceae,  377 
Melaspilea,  630 

arthonioides   (Fe"e)  Nyl.,  659,  692 
Melosira,  610,  614 

granulata  (Bhr.)  Ralfs.,  613 
Menispermaceae,  98,  388,  582 
Menispermum   canadense   I/.,  87,   98, 

131,  132,  363,  388,  582 
Mentha,  545,  628 

aquatica,  199 

canadensis  L-,  358,  363,  405,  545, 

59 r,  623,  625 

Meriolix  serrulata  (Nutt. )  Walp.,  587 
Merismopedia,  614 

glauca  Naeg.,  612 

Mesadenia  reniformis  ( Muhl. )  Raf.,  414 
Metaspermse,  373 
Micrampelis  lobata  (Michx. )  Greene, 

363.  593 
Micrasterias,  617 

truncata  (Corda)  Breb.,  612 
Micropuccinia,  543,  550 
Microsphsera,  424 

aini  (DC.)  Wint,  430 

diffusa  C.  &  P.,  430 

dubyi  Lev.,  430 

quercina  (Schw. )  Burrill,  429 

ravenelii  Berk.,  429 

russellii  Clinton,  429 

symphoricarpa  Howe,  429 
Mimulus  jamesii  T.  &  G.,  360,  406 

ringens  L.,  358,  361,  406,  591 
Mirabilis,  89 
Mitella  diphylla  L-,  363,  390 

nuda  L.,  551 
Mnium  cuspidatum  Hedw.,  43 

punctatum  Hedw.,  43 

serratum  Brid.,  43 
Mohrodendron      carolinum      (Linn. ) 

Britt.,  82 

Mollugo  verticillata  L.,  386 
Monarda  fistulosa  L.,  404,  545,  591 
Moracese,  71,  93,  383,  578 
Morus  rubra  L.,  383 
Mougeotia  parvula  Hass.  var.  angusta 

(Hass. )  Kirchn.,  26 
Muhlenbergia   racemosa   ( Michx. )   B. 

S.P.,  573 
Myrioph) Hum  spicatum  L.,  588 

verticillatum  L.,  588 


Myrtaceae,  80,  122 
Myurella  careyana  Sull.,  47 

julacea  Sch.,  47 
Myurellae,  36 

Nabalus,  425,  556 

albus  (L,. )  Hook.,  363,  409,  594 

racemosus  (Michx.)  DC.,  594 
Naiadacese,  373,  572 
Naias     flexilis      (Willd.)      Rost.      & 
Schmidt,  355,  373 

guadalupensis  (Spreng. )  Morong, 

355,  368,  373 
Napsea  dioica  L.,  397 
Napellus  vulgaris  Fourr.,  351 
Narcissus,  55,  62,  63 
Navicula,  614 
Neckera  oligocarpa  B.S.,  44 

pennata  Hedw.,  44 
Nelumbo,  643,  644,  650,  651 

lutea  (Willd.)  Pers.,  356,  357,  386, 

645,  649 
nelumbo,  649 

Nelumbium  speciosum,  645 

Nepeta  cataria  L,.,  363,  404 

Nephrocytium,  617 

agardhianum  Naeg.,  612 

Nephroma,  6,  236,  282 
epidiota  Th.  Fr.,  233 
helveticum  Ach.,  3,  14,  251 
Isevigatum  Ach.,  251,  678,  688 

var.      parile      Nyl.,     228, 

251 
tomentosum  (Hoffm.)  Koerb.,233, 

251 
Nereocystis   liitkeana,    179,    702,    726, 

732,  739 
Neurocaulon,  187 

foliosum,  176,  177,  178 
rosamarina,  176,  177 
sitchensis,  176 
Nostoc,  195,  612,  614 
lichenoides,  198 
Nostocese  Kiitz,  27 

bulliardi  Tul.,  564 
Nummularia   lateritia    Ellis   &   Ever- 

hart,  564 
nummularia    (Bulliard)  Schroet., 

564 

repanda  (Fries)  Nitscbke,  564 
Nyctaginacese,  385,  581 


INDEX. 


765 


Nymphsea,  644 

ad  vena     Soland.,    356,    386,    572, 

622,  626 

Nymphaeaccse,  386,  572,  643,  650,  651 
Nymphaeiuese,  651 

CEnothera,  427 

rhombipetala  Nutt.,  398,  587 

Oleaceae,  401 

Omphalaria,  282,  299 

kansana  Tuck.,  281,  298,  314 
phyllisca  (Wahl.)  Tuck.,  278,  280, 

29°.  3M 

pulvinata  Nyl.,  281,  298,  299,  314 
umbella  Tuck.,  299 
Onagra  biennis  (L. )  Scop.,  398,  427, 

559,  587,  627 
Onagracese,  398,  587 
Onoclea  sensibilis  L.,  358,  361,  369 
struthiopteris    (L. )    Hofftn.,    363, 

366-  369 
Onosmodium     carolinianum     (Lam.) 

DC.,  403,  590 
Opegrapha,  659,  660 

varia    (Pers. )    Fr.,   272,  303,  325, 

704 

var.  notha  Ach.,  228,  272 
var.    pulicaris    (Hoffm. )    Fr., 

281,  325 

Ophioglossaceae,  369 
Opulaster    opulifolius    (L. )     Kuntze, 

390 

Orchidaceae,  379 
Orchis  spectabilis  L.,  379 
Orobanchacese,  407 
Orthocarpus  luteus  Nutt.,  592 
Orthotrichutn,  36 

elegans  Schwaegr.,  41 
spcciosum  Nees,  41 
elegans,  41 
sroellii  Vent.,  41 
Oscillatoria,  6n,  614 

geminata  Menegh.,  28 
tenuis  Agardh.,  29 

var.  tergestina,  28 

Osmunda  claytoniana  L.,  363,  366,  369 
Osmundacese,  369 
Ostrya,  565 

virginiana      (Mill.)     Willd.,    363, 

38i 
Oxalidacese,  394,  585 


Oxalis  stricta  L-,  363,  394,  429,  585 

violacea  L. ,  367,  394 
Oxycoccus  oxycoccus  (L.)  MacM.,  589 
Oxygraphis  Bunge,  502 

Andersoni  Freyn,  499 

cymbalaria   (Pursh)   Prantl.,  427, 
503,  582 

Palmella  miniata  Leibl.   var.  aequalis 

Nag.,  26 

Palmellaceae  (Decne. )  Nag.,  26,  28 
Panax  quinquefoliuin  L.,  366,  399 
Pandoriua,  617 

morum  (Muell. )  Bory,  612 
Panicularia  americana  (Torr. )  MacM., 

375,  574 
Panicum  capillare  L.,  374,  55°.  573 

crus-galli  L.,  374,  573 

porterianum  Nash,  374 

pubescens  Lam.,  573 

scribnerianum  Nash,  21,  374 

virgatum  L.,  374,  573 
Pannaria  flabellosa  Tuck.,  253 

languinosa  (Ach.)  Kocrb.,  14,  252, 

290,  292,  298,  299,  301,  313,  680, 
689 

lepidiota  Th.  Fr.,  253 
microphylla  (Sw. )  Delis,  14,  221, 

222,  224,  253,   278,   290,  313 

nigra   (Huds.)  Nyl.,  228,  253,  298, 

299,  3'3 

petersii  Tuck.,  689 
Papaveracese,  388 
Papilionacese,  74,  392,  584 
Parietaria  pennsylvarJca  Muhl.,  384 
Parkinsonia  aculeata  Linn.,  73,  85,  87, 

102,  131,  132,  133 
Parmelia,  225,  672,  676 

borreri   Turn.,   13,   238,   272,   290, 

291,  292,  302,  310,  666,   670,  685 
var.  hypomela  Tuck.,  290,  310 
var.  rudecta  Tuck.,  245,  310 
caperata  (L. )  Ach.,  13,  222,  223, 

224,  246,  290,  292,  302,  311,  659, 

666,  670,  677,  686 
centrifuga  (L. )  Ach.,  229,  247 
cetrata  Ach.,  290,  310 
colpodes,  268 
conspersa  (Ehrh.)   Ach.,   13,  222, 

223,  224,  247,  284,  286,  288,  301, 
311,  66r,  665,  687 


766 


INDEX. 


Parmelia 

conspurcata     (Schaer. )     Wainio, 

686 
crinita  Ach.,  13,  247,  290,  292,  302, 

310,  670,  685 

encausta  (Sm.)  Nyl.,  232,  246 
olivacea  (L. )  Ach.,    13,  246,  304, 

311,  666,  670,  671,  677,  686 
var.  aspidota  Ach.,  686 
var.    prolixa   Ach.,    246,   278, 

284,  286,  311 

var.  sorediata  (Ach. )  Nyl.,  670 
perforata    (Jacq.)    Ach.,    13,    229, 

244,  302,  310 

var.  hypotropa  Nyl.,  228,  245 
perlata  (L-)  Ach.,  244 

var.  ciliata  DC.,  245 
physodes  (L. )  Ach.,  246,  675,  677, 

686 
saxatilis   (L-)    Fr.,    13,    222,    224, 

245,  246,  290,  292,  302,  310, 
658,  659,  666,  678,  685 

var.        pauniformis       (Ach.) 
Schaer.,  280,   290,   292,  310 

var.    sulcata    Nyl.,    246,    290, 

310,  670,  686 

tiliacea   (Hoffm.)   Flk.,   238,   245, 
302,  310,  670,  685 

var.    sublsevigata    Nyl.,    230, 

245 
Parnassia  caroliniana  Michx.,  361,  390, 

583 

palustris  L,.,  583 
Parnassiaceae,  583 
Parthenocissus    quinque  folia    (L.) 

Planch.,  79,  85,  358,  363,  396,  429 
Pastinaca  sativa  L.,  588 
Pediastrum  boryanum,  617 

duplex  Meyen,  611,  617 
Pedicularis  acuminatus  Dougl.,  35 

canadensis  L.,  363,  366,  406 

fluviatilis  Heller,  33 

lanceolata   Michx.,  361,  406,  425, 

592 

Pelargonium,  54 
Pellsea  atropurpurea   (L. )   Link,  367, 

371 
stelleri  (S.  G.  Gmel.)  Watt,  366, 

371 

Peltigera,  221,  225,  292,  668,  675 
aphthosa  (L. )  Hoffm.,  238,  251 


Peltigera 

canina  (L-)  Hoffm.,  14,  291,  294, 

306,  312,  667,  674,  679,  688 
var.  leucorrhiza  Flk.,  679,  689 
var.  sorediata  Schaer.,  14.  252, 
294.  3oi,  306,  312,  667,  674, 
689 

var.  spongiosa  Tuck.,  278,  312 
var.  spuria  Ach.,  14,  252,  312, 

667,  688 
horizontalis  (L. )  Hoffm.,  252,  313 

667,  688 

polydactyla  (Neck.)  Hoffm.,  252 
pulverulenta  (Tayl.)  Nyl.,  14,  252 
rufescens  (Neck.)  Hoffm.,  14,  291, 

294,  312,  688 
venosa  (L. )  Hoffm.,  251 
Penthorum  sedoides  L.,  357,  389,  582 
caudatus  Heller,  34 
gracilis  Nutt.,  591 
Peplis  portula,  199 
Peramium  pubescens  (Willd.)  MacM., 

363.  379 
Peridermium  Lev.,  537,  559 

abietinum  (A.  &  S. )  Thiim.  var. 
decoloraus  Thiim.,  559 

balsatneum  Pk.,  559 
Peridineae,  610,  617 
Peridinium,  617 

tabulatum  Ehr.,  613 
Peronospora,  196 
Pertusaria,  260,  672 

communis  DC.,   16,  260,  274,  678, 

695,  707 

finkii  A.  Zahlb.,  671,  696 
glomerata  (Ach.)  Schaer.,  229,  261 
leioplaca  (Ach.)  Schaer.,  260,  303, 

32°,  693 
multipunctata  (Turn.)  Nyl.,  260, 

683,  693 
var.  Isevigata  Turn,  and  Borr. , 

260 
pustulata    (Ach.)   Nyl.,    261,  303, 

320,  680,  696 
velata  (Turn.)  Nyl.,   16,  260,  303, 

320,  680 
Petasites    palmata    ( Ait.  )    A.    Gray, 

551 
Peucedanum  nudicaule  (Pursh)  Nutt., 

191 
Phaca  neglecta  T.  &  G.,  585 


INDEX. 


767 


Phaseolus,  55 

multiflorus,  56,  57,  58,  67 
Philotria,  54,  65,  66,  67,  68 

canadensis  (Michx.)  Britton,  360, 

374,  572 

Phleum  pratense,  622 
Phlox  divaricata  L.,  363,  402 

pilosa  L.,  402 

Phoradendron  flavescens,  169 
Phortnidium,  614 

valderiauum  (Delp. )  Goniont,  28 
Phragmidium  Link.,  537,  552,  553 

potentillae  (P.)  Karst.,  553 

rubi-idsei  (P.)  Karst.,  553 

spcciosum  Fr.,  553 

subcorticum     (Schrank.)     Wint., 

553 
Phragmites    phragmites    (L. )    Karst, 

547,  574 
Phrajjmopyxis,  552 

Phryma  leptostachya  L.,  363,  407,  559, 

592 

Phrymacese,  407,  592 
Phyllactinia,  424 

suffulta  (Reb. )  Sacc.,  429 
Phyllobium  climorphum,  198 
Phyllophora  brodiaei,  163 

interrupta,   163 
Physalis  heterophylla  Nees,  405 

macrophysa  Rydb.,  405 

philadelphica  Lam.,  358,  405 

virginiana  Mill.,  405 
Physcia,  222,  224,  672,  676 

adglutinata   (Floerk.)    Nyl.,    228, 

249,  303,  3°4,  312,  688 
aquila  (Ach. )  Nyl.,  230,  247 

var.  detonsa  Tuck.,  14 
astroidea  (Fr. )  Nyl.,  666,  687 
csesia  (Hoffm.)  Nyl.,  14,  224,  228, 

249,   284,  287,  288,  312,  661,  664, 

688 

ciliaris  (L.)  DC.,  247 
granulifera  (Ach.)  Tuck.,  238,  303, 

311,  687 
hispida  (Schreb.,  Fr.)  Tuck.,  227, 

248,  666,  671,  675,  677,  687 
hypoleuca    (Muhl.)    Tuck.,    670, 

687 
obscura  (Ehrh.)  Nyl.,  14,  222,  223, 

248,  249,  290,  292,  303,  312,  666, 

671,  680,  688 


Physcia 

pulvcrulenta    (Schreb.)  Nyl.,    14, 

247,  249,  290,  292,  303,  311, 
666,  671,  687 

var.  leucoleiptes  Tuck.,  248 
speciosa   (Wulf.,   Ach.)   Nyl.,  13, 

222,  247,  290,  292,  311,  666,  670, 

687 
stellaris   (L. )   Tuck.,   14,  222,  224, 

248,  292,  303,  311,   659,666, 
687 

var.  apiola  Nyl.,  248,  287,  288, 

311,  664,  666,  687 
tribacia  (Ach. )  Tuck.,  14,  248,  284, 

287,  303,  312,  681,  687 
Physostegia    virginiana    (L. )  Benth., 

358,  404,  591 
Picea  mariana    (Mill.)     B.S.P.,     225, 

559 
Pimpinella  integerrima  (L. )  A.  Gray, 

361,  399 
Pinacese,  372 
Pinus  Banksiana,  32 
coronaria,  31 
ponderosa,  538 

scopulorum,  169 
resinosus  Ait.,  32,  225 
strobus  L,.,  225,  372 
Pistia,  649 
Placodium,  221,  665,  672,  674 

aurantiacum    (Light)    Naeg.    and 
Hepp.,    15,    223,    255,   291,   300, 
3°3,  3 '5,  664,  671,  690 
cerinum      (Hedw. )      Naeg.      and 
Hepp., 255,303.  304,  316,  664, 
671,  691 
var.   pyracea  Nyl.,    229,    255, 

305,  316,  673,  691 
var.  sideritis  Tuck.,   15,  285, 

287,  288,  316,  664,  691 
cinnabarinum    (Ach.)    Auz.,    15, 
228,  255,  285,  287,  297,  315,  664, 
690 
citrinum   (Hoffm.)    Leight.,   255, 

298,  299,  316 
elegans  (Link.)  DC.,   15,  255,  284, 

287,  300,  315,  661,  664,  690 
ferrugineum    (Huds. )    Hepp., 

281,  316,  673,  691 
var.      pollinii     Tuck.,      281, 


768 


INDEX. 


Plac  odium 

murorum   (Hoffm.)  DC.,  229,  255, 

287,  315,  651,  690 
var.  nriniatum  Tuck.,  228,  255 
vitellinum     (Ehrh.)     Naeg.     and 
Hepp.,  15,  223,  224,  229,  256, 
284,  287,  289,  316,  664,  691 
var.  aurellum  Ach.,  256,  2971 

300,  316,  669,  691 
Plagiochila     asplenioides     (L. )      Du- 

mort,  193 

Plagiothecium  denticulatum  Sch.,  50 
muhlenbeckii  Sch.,  50 
sylvaticum  Sch.,  50 
Plantaginacese,  407 
Plantago  eriopoda  Torr.,  571,  592 

major  L.,  407,  592 
Purshii,  22,  23 
Platygyrium  repens  Sch.,  48 
Pleurotseniopsis,  617 

quaternaria    (Nordst. )    De   Toni., 

612 
Poa  pratensis  L.,  428,  622 

pseudopratensis    Scrib.    &    Ryd., 

192 

Podophyllum  peltatum  L.,  363,  388,  549 
Podosphsera,  424 

oxyacanthae  (DC.)  D.  By.,  429 
Pogonatum  alpinum  Roell.,  43 
Polanisia  graveoleus  Raf.,  389 
Polemoniaceae,  402 
Polemonium  reptans  L-,  363,  402 
Polycystis,  28 
Poly  gala  senega  L.,  395 
verticillata  L.,  367,  394 
viridescens  L.,  394 
Polygalaceae,  394 
Polygonacese,  145,  384,  578 
Polygonatum,  554 

commutatum    (R.  &    S. )    Dietr., 

363,  366,  378,  577 
commutatum  (Sch.)  Dietr.,  554 
Polygonum,    142,    540,   542,   543,    622, 

628 

amphibium  I,.,  549 
aviculare  L.,  427,  542 
convolvulus  L.,  385,  579 
divaricatum,     138,    139,    141,    142, 

145,  146,  149 

emersum    (Michx. )    Britton,    356 
357,  384,  549,  579 


Polygonum 

erectum  L.,  137,  142,  143,  144,  146, 

151,  579 

exsertum  Small,  579 
hartwrightii    A.    Gray,    358,    384, 

549 

hydropiper  L.,  384,  627 
hydropiperoides,  358,  363 
incarnatum  Ell.,  358,  363,  384,625 
lapathifolium  L.,  579 
littorale  Link,  579 
orientale  L.,  384 
persicaria  L.,  579 
punctatum  Ell.,  358,  384,  579 
ramosissimum    Michx.,    385,   542, 

579 

sagittatum  I/.,  385 
scandens  L.,  385,  579,  627 
tenue  Michx.,  367,  368,  385 
virginianum  L.,  358,  368,  385 
Polyides  lumbricalis,  195 
Polymnia  canadensis  L-,  412 
Polypodiaceae,  369 
Polypodium  vulgare  L.,  371 
Polyporus  versicolor,  694 
Polysiphonia,  197,  199,  203 
Polytrichum  commune  I/.,  43 
juniperinum  Willd.,  43 
piliferum  Schreb. ,  43 
Pomaceae,  391,  584 
Populus,  254,  428,  539,  564,  567 
alba  L.,  380 
balsamifera  I/.,  225,  577 

candicans     (Ait.)     A.     Gray, 

380 
deltoides  Marsh,  69,  358,  380/428, 

540,  624,  626,  629 
grandidentata    Michx.,    363,    380, 

428 
tremuloides  Michx.,  225,  363,  380, 

428,  539,  624,  626,  629 
Porella  platyphylla   (L,. )    L/indb.,  193 
Porphyra  pertusa,  206 
Potamogeton  lonchites  Tuckerm.,  356, 

373 

natans  L.,  356,  373 
pectinatus  L.,  572 
perfoliatus  L.,  572 
pusillus  L.,  355,  373 
zostersefolius  Sch u in.,  355,  373 
Potamogetoninae,  651 


INDEX. 


769 


Potentilla  arguta  Pursh,  390 
canadensis  L.,  31,  363,  391 
effusa  Doug].,  569   583 
hippiaua  Lehtn.,  191 
leucocarpa  Rydberg,  30,  583 
monspeliensis    L.,    30,    390,    583, 

626 

Nicolletii  (Wats.)  Sheldon,  30,  31 
pennsylvauica  strigosa  Pursh,  553, 

583 

pentandra  Engeltn.,  31 

supitia  var.  Nicolletii  Wats.,  31 
Prenanthcs,  546 
Pritnulaceae,  400,  589 
Protococcaceie,  198 
Protococcus,  195,  617 
Protomyces,  199 
Prunella  vulgaris  L-,  363,  404 
Prunus,  429 

amcricana  Marsh,  363,  392,  584 

nigra  Ait.,  363,  368,  392 

pumila  I,.,  549 

serotina  Ehrh.,  363,  392,  584 

virginiana  L  ,  363,  366,  392 
Psoralea  argophylla  Pursh,  543,  584 
Ptelea  trifoliata  Linn.,  75,  84,  87,  in, 

130,  131,  132,  133 
Pteridophyta,  369 
Pteris  aquiliua  L.,  363,  366,  371 
Pterygophora  californica  Ruprecht,  723 
Ptilidium  ciliare  (L. )  Nees,  193 
Puccinellia  airoides   (Nutt. )   Wats.  & 

Coult.,  569,  574 
Puccinia  Persoon,  537,  543,  545,  551,552 

adoxse  Hedw.,  544 

aegra  Grove,  640 

alpina  Fckl.,  640 

amorph(Z  Curt,  544,  552 

anemones-virginianae,  537 

angustata  Peck,  546 

argentata  (Schultz)  Wiut.,  549 

asteris  Duby,  550 

calthse  Lk.,  544 

caricis  (Schuui.)  Rebent,  546 

Chondrillse  Corda,  546 

circiese  Pers. ,  550 

convolvuli  (P.)  Cast.,  544 

deglubens,  552 

densa  D.  &  H.,  633,  639 

effusa  D.  &  H.,  639 

elytni  \Yestd.,  552 


Puccinia 

emaculata  Schwcin.,  550 
Fergussoni  Berk.  &  Br.,  639,  640 
Fergussoni  hastattr  DeT.,  633 
galii  (P.)  Schwein.,  544 
gentians  (Strauss)  L,k.,  544 
graininis  Pers.,  548 
grossulariae  (Gm.)  Wint.,  541 
halenire  Arth.  &  Holway,  551 
hastatcs  Cooke,  633,  637,  638 
hieracii  (Schutn. )   Mart.,  543,  548 
hydrophylli  Peck  &  Clint.,  551 
kuhniae  Schwein.,  549 
Mariae-Wilsoni  Clinton,  639 
menthse  americana  Burrill,  545 
mesomegala  B.  &  C.,  551 
nardosmiae  E.  &  E.,  551 
ornata  Arth.  &  Holw.,  550 
peckiana  Howe,  552 
petalostemonis  Farlow,  552 
phragmitis  (Schum. )  Korn,  547 
pimpinellae  (Strauss)  Link,  544 
poculiformis  (Jacq. )  Wett.,  548 
podophylli  Schwein.,  549 
polygoni-amphibii  P.,  549 
porphyrogenita  Curt.,  551 
pruni-spinosae  Pers.,  549 
rhamni  (P.)  Wettst.,  547 
rubigo-vera  (DC.)  Wint.,  548 
Schceleriana  Plow,  et  Magn.,  556 
sorghi  Schwein.,  550 
tanaceti  DC.,  545 
taraxaci  Plowright,  548 
thalictri  Chev.,  550 
tiarellae  B.  &  C.,  551 
totnipara  Trel.,  549 
variolans  Hark.,  550 
violae  (Schum.)  DC.,  544,  631,  633, 

640 

Violarum  Link.,  633 
xanthii  Schwein.,  551 

Pucciniaceae,  537,  540 

Pucciniopsis,  543,  549 

Pulsatilla  hirsutissima(  Pursh)  Britton, 

367,  387 

Pylaisia  heteromalla  Sch.,  48 
polyantha  Sch.,  48 

Pylaisiellse,  36 

Pyrenopsis,  282 

melambola  Tuck.,  280,  313 
phaeococca  Trek.,  280,  313 


770 


INDEX. 


Pyrenula,  225,  227,  305 

cinerella  (Plot.)  Tuck.,  228,  275, 

304,  328,  709 

var.      quadriloculata     Fink., 
228,    276,    304,     329,      672, 
709 
gemmata  (Ach.)  Naeg.,  281,  303, 

328,  709 

glabrata  (Ach.)  Mass.,  281,  329 
hyalospora  (Nyl. )  Tuck.,  281,  303, 

328,  709 
leucoplaca     (Wahl.)     Kbr.,     225, 

239.  275,  3°3.  3°4,  329>  67i, 
672,  709 

var.    pluriloculata  Fink,   671, 

672,  709 

megalospora  Fink,  281,  303,  329 
nitida  Ach.,  303,  304,  328 
punctiformis   (Ach.)    Naeg.,   304, 

328,  708 
var.  fallax  Nyl.,  275,  276,  304, 

328,  672,  709 
quinqueseptata  (Nyl.)  Tuck.,  281, 

303,  329 

thelena  Ach.,  304,  328 
Pyrola  elliptica  Nutt.,  363,  400,  538 

rotundifolia  L.,  538 

secunda  L.,  538 
Pyrolaceae,  400 

Pyrrhopappus  Rothrockii  A.  Gray,  35 
Pyrus  coronaria  var.  loensis  Wood,  31 

loensis  Bailey,  31 
Pyxine  sorediata  Fr.,  14,  249,  290,  312 

Quercus,  564,  565,  566 
alba  L.,  366,  382,  578 
coccinea  Wang.,  363,  365,  382 
macrocarpa  Michx.,  239,  363,  365, 

366,  382,  429,  578 
platanoides    (Lam. )   Sudw.,    358, 

368,  382 

prinoides  Willd.,  368,  383 
rubra,  363,  365 
velutina  Lam.,  363,  382 

Radula  complanata  (L.)  Dumort.,  193 

Ramalina,  225,  236,  672 
calicaris  (L.)  Fr.,  291 

var.  canaliculata  Fr.,  230,  240 
var.    farinacea     Schaer.,     13, 

240,  290,  301,  308,  309 


Ramalina 

calicaris  (L,.)   Fr.,  var.    fastigiata 

Fr.,  229,  240,  302, 308,670, 683 

var.    fraxinea   Fr.,    302,    308, 

670,  683 

pollinarella  Nyl.,  229,  241 
pollinaria  (Ach.)  Tuck.,  241 
polymorpha    (Ach.)    Tuck.,    280, 

290,  308 
pusilla    (Prev.)    Tuck.,   230,    241, 

662,  677,  678,  683 
var.  geniculata  Tuck.,  232,  241 
Ranunculaceae,  89,  387,  581 
Ranunculus,  459,  462,  479,  500 

abortivus  Hook.,  363,  387,  477,  478, 

479,  555 

var.  australis  Brand.,  478 
var.    grandiflorus     Eugelm., 

478 

var.  Harveyi  Gray,  478 
var.  micranthus  Gray,  470 
aconitifolius  Linn.,  492 
acriforniis  Gray,  476 
acris  Hook.,  476 
acris  Linn.,  471,  475,  481 
acris  var.  Deppii  Nutt.,  476 
adoneus  Gray,  488 
affinisR.  Br.,  482 
affinis  Torr.,  484 

var.  cardiophyllus  Gray,  482 
var.  lasiocarpus  Torr.,  482 
var.   leiocarpus  Traut,  482 
var.  micropetalus  Greene,  483 
var.  validus  Gray,  482 
alceus  Greene,  468 
Allegheniensis  Britton,  478 
alpeophilus  A.  Nelson,  483 
alismcsfolius  Benth.,  494 
alistnsefolius  Geyer,  495 

var.  alismellus  Gray,  495 
var.  calthseflorus  Davis,  495 
alismaffoliusv&r.  montanus  Wats., 

495 
alismellus  Greene,  495 

var.  populago  Davis,  496 
ambigens  Wats.,  494 

var.  obtusiusculus  Davis,  494 
amarillo  Bertol.,  475 
amcenus  Gray,  488 
amcenus  Ledeb.,  482 
atnplexicaulis  Linn.,  493 


INDEX. 


771 


Ranunculus 

Anderson!  Gray,  499 

Andersons  var.  tend! us  Wats. ,499 

aquatilis  Linn.,  460,  461 

var.  brachypus  Hook.  &  Arn., 

461 

var.  cirspitosus  DC.,  461 
var.  confervoides  Gray,  461 
var.  (titan  cat  us  Gray,  460 
var.  flaccid  us  Gray,  461 
var.  hetcrophyllus  DC.,  461 
var.  kispidvlus  Drew,  461 
var.  Lobbii  Wats.,  462 
var.  longirostris  Lawson,  460 
var.  stagHatalis  DC.,  460 
var.  submersiis  Gordon,  461 
var.  trichophyllus  Gray,  461 

arcticits  Richards.,  482 

arcualus  Heller,  480 

Arizonicus  Greene,  485 

var.  subaffinis  Greene,  483 

Arizonicus  Lemmon,  484 

arnoglossus  Greene,  494 

arvensis  Linn.,  467 

Aschenbornianus  Schau.,  476 

Asiaticus  Linn.,  477 

auricoinus  Hook.,  482 

var.  Cassubicus  E.  Meyer,  484 

Austin  a:  Greene,  489 

Beckii  G.  Don,  491 

Belvisii  DC. ,  470 

Biolettii  Greene,  499 

Bloomeri  Wnts.,  472 

Bolanderi  Greene,  495 

Bonariensis  Poir. ,  499 

Bongardi,  Greene,  479 

var.Douglasii  (Ho well)  Davis, 

479 
var.  tenellus  Greene,  480 

breincaulis  Hook.,  484,  489 

bulbosus  Linn.,  471,  472 

Caleottii  Turcz.,  468 

Californicus  Benth.,  476 
var.  canus  Wats.,  474 
var.  crassifolius  Greene,  477 
var.  latilobus  Gray,  477 
var.    Ludovicianus    (Greene) 
Davis,  494 

calthceflorus  Greene,  495 

Canadensis  Jacq.,  469 

canus  Benth.,  474 


Ranunculus 

cardiophyllus  Hook. ,  482 

var.  pinetorum  Greene,  483 

carpaticus  Herbich.,  485 

Chilensis  Hook.  &  Arn.,  472 

ciliosus  Howell,  481 

drcinatus  Sibth.,  460 

Clintonii  Beck,  475 

confervoides  Fries.,  461 

Cooleyte  Vasey  &  Rose,  502 

corthusaefolius  Willd.,  486 

Cusickii  Jones,  496 

Cymbalaria  Pursh,  503 

var.  alpinus  Hook.,  503 

delphinifolius  H.B.K.,  474 

delphinifolius  Torn,  387,  491 

delphinifolius  Torr.  &  Gray,  476 

dichotonms  Moc.  &  Sesse,  473 

digitatus  Hook.,  489 

dissectus  Hook.  &  Arn.,  476 

divaricatus  Schrank,  460 

Donianus  Pritz  ,  486 

Douglasii  Howell,  480 

Drummondi  Greene,  488 

Earlei  Greene,  479 

Eiseni  Kellogg,  480 

ellipticus  Greene,  490 

eremogenes  Greene,  478 

var.  degeuer  Greene,  478 

Eschscholtzii  Schlecht.,  483 

eximius  Greene,  483 

fascicularis  Brittou,  472 

fascicularis  Muhl.,  470 

fascicularis  Schlecht.,  471 

fascicularis  Wats.,  469 

fascicularis   var.   Deforesti  Davis, 
470 

Ficaria  Linn.,  501 

filifortnis  Michx.,  497 

flaccidus  Pers.,  461 

Flammula  Hook.,  495 

Flammula  Michx.,  496 

Flammula  Pursh,  494 

Flammula  Walt.,  499 

var.  filiform  is  Hook.,  497 
var.  intermcdiusHoob..,  498 
var.  laxicaulis  Torr.  &  Gray, 

496 

var.  reptaus  E.  Meyer,  497 
var.  Unalaschensis  Ledeb.,498 

flaviatilis  Bigel.,  491 


772 


INDEX. 


Ranunculus 

geoides  H.B.K.,  487 
glaberrimus  Hook.,  489 
glaberriraus  var.  ellipticus  Greene, 

490 

glabriusculus  Rupr.,  480 
glacialis  Linu.,  500 
Gmelini  DC.,  492 
Gotmani  Greene,  500 
Grayanus  Freyn,  461 
Grayi  Britton,  488 
Greenei  Howcll,  479 
halophilus  Schlecht,  503 
Hartwegi  Greene,  495 
Harveyi  Britton,  478 
h'ebecarpus  Hook.  &  Arn.,  468 
hebecarpus  var.  pusillus  Wats.,  468 
hederaceus  Linn.,  462 
,         var.  Torr.   462 

var.  Lobbii  Lawson,  462 
hesperoxys  Greene,  475 
hirsutus  Curt.,  469 
hispidus  Hook.,  469 
hispidus  Michx.,  472 
hispidus  Pursh,  469 

var.  Oreganus  Gray,  469 
Hookeri  Schlecht.,  471 
Hooker i  Reo el,  488 
Howellii  Greene,  480 
humilis  D.  Don,  486 
humilis  Pers.,  499 
hydrocharus  subsp.  Lobbii  Hiern, 

462 

hydrocharoides  Gray,  496 
hyperboreus  Rottb.,  491 

var.  natans  Regel,  491 
hystriculus  Gray,  500 
Icelandicus  Davis,  472 
inamcenus  Greene,  483 
intermedius  Heller,  498 
juuiperiuus  Jones,  499 
lacustris  Beck  &  Tracy,  491 
Lambertianus  D.,  493 
Langsdorfii  DC.,  492 
lanuginosus  Walt.,  479 
Lappotiicus  Linn.,  491 
Lapponicus  Oed.,  489 
laxi caulis  Darby,  496 
Ivrnmoni  Gray,  496 
limosus  Nutt.,  492 
lingua  Pursh,  494 


Ranunculus 

Llavseuus  Schlec-ht.,  473 
Lobbii  Gray,  462 
longipedunculatus  Scheidw.,  486 
longirostris  Godron,  460 
lucidus  Poir.,  470 
Ludovicianus  Greene,  476 
Lyalli  Rydb. ,  479 
Macauleyi  Gray,  490 
Macounii  Britton,  469 

var.  Oreganus  Davis,  469 
macranthus  Scheele,  474 
macranthus  Wats.,  473 
Madrensis  Rose,  495 
Marilandicus  Poir,  472 
maximus  Greene,  473 
McCallai  Davis,  482 
Mexicanus  Davis,  487 
micranthus  Nult.,  470 
microlonchus  Greene,  498 
Missouriensis  Greene,  492 
Montanensis  Rydb.,  476 
montanus  Willd.,  485 
montanus  var.   dentatus  Baumg., 

485 

var.  subaffinis  Gray,  485 
var.  subsagittatus  Gray,  485 

tnulticaulis  D.  Don,  486 

multifidus  Pursh,  491 

var.   limosus  Lawson,  492 
var.  repens  Wats.,  492 
var.  terrestris  Gray,  491 

muricatus  Linn.,  467 

natans  C.  A.  Meyer,  490 

Nelsoni  Gray,  480 

var.  giabriusculus  Holzinger, 

480 
var.  tenellus  Gray,  479 

nilidus  Walt.,  477 

nivalis  Linn.,  490 

nivalis  Rep.,  490 

nivalis   var.  Eschscholtzii  Wats., 

483 

Nuttallii  Gray,  502 

oblongifolius  Ell.,  496 

obtusiusculus  Biitton,  494 

obtusiusculus  Raf.,  494 

occidentalis  Gray,  481 

occidentalis  Nutt.,  480 

var.  brevistylus  Greene,  480 
var.  Eiseni  Gray,  480 


INDEX. 


773 


Ranunculus 

var.  Howellii  Greene,  480 
var.  Lyalli  Gray,  479 
var.  parviflorus  Torr.,  479 
var.  Rattani  Gray,  481 
var.  robustus  Gray,  481 
var.  ten  el  I  us  Gray,  480 
var.    ultramontanus    Greene, 
481 

Of  real  us  Greene,  483 

Oreganus  Ho  well,  469 

ornithorhynchus  Walp.,  473 

orthorhynchus  Hook.,  473 
var.  alpinus  Wats.,  488 
var.  platyphyllus  Gray,  473 

ovalis  Raf.,  484 

oxynotus  Gray,  489 

Pallasii  Schlecht. ,  493 

palmatus  Ell.,  475 

pantothrix  Brot.  ex  DC.,  461 

parviflorus  Linn.,  468 

parviflorus  var.  Torr.  &  Gray,  468 

parvuliis  Linn.,  469 

pedalifidus  Hook.,  488 

pedatifidus  J.  E.  Smith,  482 

var.  cardiophyllus  Britton,  482 
var.       pinetorum       (Greene) 
Davis,  483 

pennsylvauicus    Linn.,    358,  387, 
469,  581,  627 

petiolaris  H.B.K.,  487 

Philonotis  Ebrh.,  469 

Philonotis  Pursh,  470 

pilosus  H.B.K.,  471 

Populago  Greene,  496 

Porteri  Britton,  461 

prostratus  Poir.,  475 

Purshii  Richards,  492 

Pursh ii  Torr.,  490 

Purshii  var.  aquatilis  Ledeb.,  491 

pusilltis  Ledeb.,  492 

pusillus  Poir.,  499 

var.  Lindheimeri  Gray,  499 
var.     oblongifolius    Torr.    & 
Gray,  456 

pygmseus  Wahl.,  489 

var.  Sabinii  Davis,  489 

tadicans  C.  A.  Meyer,  490 

radieans  Regel,  492 

ranunculi HUS  Rydberg,  502 

Rattani  Howell,  481 


Ranunculus 

recurvatus  Bong.,  479 

recurvatus  Poir.,  479 

recurvatus  Schlecht.,  480 
var.  h'elsonii  DC.,  480 

regulosus  Greene,  476 

repeus,  471 

repens  Linn.,  475 

var.  hcsperoxys  Davis,  475 
var.    liispidus   Torr.   &   Gray, 

469 

var.  inacranlhus  Gray,  474 
var.    Marilandicus     Torr.    & 
Gray,  472 

rcptans  Linn.,  497 

var.  filiform  is  DC.,  497 
var.  Gormani  Davis,  498 
var.   intcrmedius  Torr.  & 
Gray,  498 

rep  tans   strigulosus   var.    Freyn.> 
498 

rhomboides  Goldie,  484 

Sabinii  "Si.  Br.,  489 

saluginosiis  Pall,  503 

samolifolius  Greene,  497 

sanicula-forinis  Muhl.,  479 

5-ardous  Crantz,  469 

saxicola  Rjdb.,  484 

scleratns  L.,  478,  581 

var.  multifidus  Nutt.,  478 

Schlechtendalii  Hook.,  471,  480 

septentrionalis  Poir.,  363,  388,470, 

475 
var.  Blankinshipii   Robinson, 

475 

speciosus  Hort.,  472 
stagnatalis  Wallr.,  460 
stolonifer  Hemsl.,  497 
subaffinis  Rydb.,  485 
subalpinus  Davis,  474 
subsagittatus       var.        subaffinis 

Greene,  485 
Suksdorfii  Gray,  484 
suphureus  Soland.,  490 
ienellus  Nutt.,  479 
tenellus    var.    Lyallii    Robinson, 

479 

tomentosus  Poir.,  470 
tomentosus  Spreng.,  479 
trachy sperm  us  Ell.,  468 
tiacbyspermus  Engelm.,  498 


774 


INDEX. 


Ranunculus 

var.    angustifolius    Engelm., 

498 
var.    Lindheimeri     Engelm., 

499 

trachophyllus  Chaix.,  461 
tridentatus  H.B.K.,  503 
trifolius  Moench.,  469 
triternatus  Gray,  488 
Turner!  Greene,  481 
Unalaschensis  Bess.,  498 
uncinatus  D.  Don,  487 
unguiculatus  Greene,  494 
vagans  Wats.,  497 
vicinalis  Greene,  483 
Raphidium,  28,  6n 

polymorphum  Fresen.    var.    acic- 
ulare  (A.  Br. )  Rabenh.,  611, 
617 
var.       falcatum        (  C  o  r  d  a  ) 

Rabenh.,  612,  617 
var.  sigmoidea,  617 
Raphidostegium  recurvans  L-   and  J., 

50 
Ratibida  columnaris  (Sims)  D.    Don, 

367,  413.   597 

pinnata  (Vent.)  Barnhart,  413 
Razoumofskya,  170 

robusta    (Engelm.)    Kuntze,   169, 

173 

Rhabdoweisia  denticulata  B.  S.,  37 
Rhamnacese,  78,  116,  396 
Rhamnus,  119 

alnifolia  L'Her.,  548 

purshiana    DC.,    78,    85,   87,   118, 

130,  131,  132,  133 
Rhodomenia,  205,  206 
Rhodymenia,  206 

palmata,  206 

pertusa,  206 

peruviana,  206 
Rhns,  566 

glabra  L-,  363,  367,  395-  5§6 

hirta  (L.)  Sudw.,  395 

radicans  L.,  358,  367,  395,  543 
Ribes,  558 

cynosbati  L.,  363,  390,  558 

floridum  L'Her.,  363,  390,  425,  558 

gracile  Michx.,  558 

rubrum  L,.,  549 

uva-crispa  L,.,  363,  390 


Ricinus,  88 
Rinodina,  665,  662 

ascociscana  Tuck.,  259 

bischoffi    (Hepp.)  Kbr.,   281,  297 

320,  669,  695 
lecanorina   Mass.,    281,    282,    287, 

288,  320 
nigra,  695 

oreina  (Ach.)  Mass.,  224,  259,  278, 
285,  287,  289,  319,  660,  664,  672, 
6?3»  694 

sophodes  (Ach.)  Nyl.,  16,  259, 
287,  288,  289.  304,  305,  319, 
659,  664 

var.  atrocinerea  Nyl.,  695 
var.  confragosa  Nyl.,  230,  259 
var.'exigua  Fr.,  305,  320,  664, 

695 

var.  tephraspisTuck.,  280,  320 
Rivularia  biasolettiana  Menegh.,  27 
Rivulariaceae  Rabenhorst,  27 
Robinia  pseudacacia  Linn.,  75,  84,  87, 

109,  130,  131,  132,  133 
Rodhymenia,  206 

Roripa  hispida  (Desv. )  Britton,  582 
nasturtium  (L,. )  Rusby,  360,  389 
palustris  (L. )  Bess.,  389,  582 
Rosa,  553,  554 

acicularis  Lindl.,  554 
arkansana  Porter,  20,  21,  391,  584 
blanda  Ait,  391,  554 
Rosacese,  390,  583 
Rubiacese,  83,  89,  130,  407,  592 
Rubus,  552 

canadensis  L,.,  390,  552 
hispidus  L-,  424,  553 
occidentalis  L,.,  363,  366,  390 
strigosus  Michx.,  552,  553,  583 
villosus  Ait.,  363,  366,  390,  552 
Rudbeckia  hirta  L/.,  412,  597,  627 

laciniata   L.,    361,    363,    413,    426, 

543,  597 

triloba  L.,  361,  363,  368,  412 
Rutnex,  138,  146,  550,  627 

acetosella  L.,  137,  361,  384 
britannica  I/.,  547,  550 
crispus  L,.,  361,  384,  547,  579 
hydrolapathum,  547 
obtusifolius,  199,  547 
occidentalis  S.  Wats.,  569,  578 
persicarioides  L,.,  579 


INDEX. 


775 


Rumex 

salicifolius  Weinm.,  137,  142,  143, 

144,  145,  148 
verticillatus,    137,    143,    144,    145, 

146,  147,  148,  578 
Ruppia  occidentalis,  570 
Rutacese,  75,  in,  394,  585 

Sagedia,  236 

oxyspora    (Nyl.)   Tuck.,  225,  230 

275,  660 

Sagina  nodosa  (L. )  Fenzl.,  620 
Sagittaria,  355,  357 

arifolia  Nutt.,  572 

cristata  Engelm.,  374 

cuneata  Sheldon,  356,  369,  374 

latifolia  Willd.,  356,  357,  374,  572, 
624 

rigida  Pursh,  356,  374 

variabilis,  144 
Salicaceae,  69,  380,  577 
Salicornia,  570,  571 

herbacca  L,. ,  580 
Salix,  428,  539,  540,  626 

amygdaloides   Anders.,    358,    363, 
380,  624  625,  629,  630 

bebbiaua  Sarg.,  381,  428 

cordata,  626 

discolor  Muhl.,  381,  428,  540 

fluviatilis  Nutt.,  358,  363,  381 

humilis  Marsh,  381 

lucida  Muhl.,  380,  578 

uiyrtilloides  L.,  428,  540 

uigra  Marsh.,  358,  380 
Salsola  tragus  L-,  385 
Salvinia  natans,  66,  67,  68 
Salviniaceit,  371 
Sainbucus,  84 

canadensis  L.,  363,  407 

pubens  Michx.,  83,  407 
Sanguinaria  canadensis  L.,  388,  399 
Sanguisorba  DC.,  35 
Sanicula   gregaria  Bicknell,  363,  369, 

399 

marylandica  L.,  363,  399 

trifoliata  Bicknell,  368,  399 
Santalaceae,  384 
Sarcophyllis,  202 

arctica,  198,  199 

edulis,  199 
Saxifraga  penns) Ivanica  L.,  361,  389 


Saxifragaceae,  389 

Scapatiia  glaucocephala  (Tayl.)'Aust., 

193 
Scenedesmus,  28,  6n 

bijugatus  (Turp.)  Kg.,  611,  617 
obliquus  (Turp.)  Kg.,  26,  617 
quadricauda    (Turp.)    Bre"b.,   611, 

617 

Schedonnardus  paniculatus,  22 
Scheuchzeriaceae,  572 
Schinus    molle  Linn.,  76,  84,  85,  87, 

114.  131 

Schizonema,  197 
Schizosporacae,  537 
Schizothrix  lardacea ( Cesati )  Gomout. , 

28 

rupicola  Tilden,  28 
Scirpus atrovirens  Muhl.,  357,  376,  546, 

575 

campestris  Britton,  569,  575 

cyperinus  (L.)  Kunth.,  357,  376 

lacustris  L-,  356,  376,  575 
Scotinosphaera,  198 
Scrophularia  marylandica  L,.,  405 
Scrophulariacese,  405,  591 
Scutellaria  cordifolia  Muhl.,  404 

galericulata  L,.,  591,  626 

lateriflora  L.,  358,   404,   590,  623, 
625 

parvula, Michx.,  404 
Selaginella  rupestris  (L. )  Spring.,  372 

selaginoides  (L. )  Link.,  619 
Selaginellaceae,  372 
Senecio  aureus  L-,  145,  415,  426,  556 

discoideus  (Hook.)  Britton,  620 

plattensis  Nutt.,  368,  414 

vulgaris  L.,  192 
Sileue  alba  Muhl.,  361,  363,  386 

antirrhina  L.,   137,   142,  367,  386, 
58i 

stellata  (L.)  Ait.,  366,  386 
Silphium  laciniatum  L.,  361,  412 

perfoliatum  L.,  361,  412 
Simarubaceae,  76,  112 
Sisymbrium  altissimum   L.,   191,  388, 
627 

canescens  Nutt.,  30 

humile  Meyer,  620 

multifidum,  30 

officinale  (L.)  Scop.,  388 
Sisyrinchium  angustifolium  Mill.  379, 


776 


INDEX. 


Sitilias  multicaulis   (DC.)  Greene,  35 

Rothrockii  (A.   Gray)   Greene,  35 
Sium   cicutsefolium    Gmel.,  357,    399, 

588 

Smilaceae,  378,  577 
Smilax  herbacea  L-,  363,  366,  378,  577 

hispida  Muhl.,  366,  379 
Solanaceae,  405,  591 
Solauum  nigrum  L.,  405,  591,  622,  627 
Solidago  canadensis  L.,  363,  411,425, 
539,  595,  627 

erecta  Pursh,  369,  410 

flexicaulis  L.,  41°,  539,  557 

hispida  Muhl.,  410 

missouriensis  Nutt.,  411 

mollis,  22,  23 

nemoralis  Ait.,  411,  596 

rigida  L.,  20,  21,  22,  411,  595 

rigidiuscula,  21 

rupestris,  20,  21,  22 

serotina  Ait.,  410,  539,  557,  627 

speciosa  Nutt.,  410 

ulmifolia  Muhl.,  410 
Solorina,  236,  662 

saccata  (L. )  Ach.,  227,  252 
Soiichus  arvensis  L.,  593 

asper  (I..)  EH.,  409,  593 
Sophia  hartwegiana  (Fourn.)  Greene, 
30,  582 

incisa,  30 

pinnata  (Walt.)  Britton,  30 
Sorghum,  550 
Sparganiacese,  373,  571 
Sparganium  eury  carpum  Engelm. ,  357, 

373,  571 

simplex  Huds.,  571 
Spartina  cynosuroides  (L.)  Willd.,  357, 

375,  574 

Spermatophyta,  372 
Sphacelaria,  199 
Sphaerococcus,  205 
Sphaerotheca,  424 

castagnei  Lev.,  424 

humuli  (DC.)  Burrell,  424 

mors-uvae  (Schw. )  B.  &  C.,  425 

pannosa  Lev.,  425 
Sphagnum,  26,  199 

acutifolium  Ehrh.,  36 

fuscum  Klinggr.,  36 

girgensohnii  Russ.,  36 

medium  Limpr.,  36,  37 


Sphagnum 

recurvum  parvifolium  Sondtn.,  37 

squarrosum  Pers.,  37 

teres    squarrosulum     (Lesq.) 

Warnst.,  37 
wulfianum  Girg.,  37 
Spiraea  salicifolia  L.,  390,  583 
Spirodela  polyrhiza  Schleid.,  355,  377 
Spirogyra,  617 
Sporobolus,  21,  23 

brevifolius  (Nutt.)  Scribn.,  573 
cuspidatus  (Torr.)  Wood,  573 
heterolepis  A.  Gray,  573 
Stachys  palustris  L.,  358,  404,  428,  591 
Staphylea  trifolia  L.,  363,  395 
Staphyleaceae,  395 
Staurastrum  crenulatum,  617 

minneapoliense  Wolle,  612,  617 
paradoxum  Meyen.  var.  longipes 

Nordst,  612 

sebaldi  Reinsch.,  612,  617 
Staurothele  diffractella  (Nyl.)  Tuck., 

281,  290,  297,  300,  327 
drummondii  Tuck.,  228,  275,  785, 

290,  328,  708 
umbriua  (Wahl.)  Tuck.,  274,  285, 

290,  298,  299,  327 
var.  colpima  (Whubl.)   Nyl., 

708 
Steironema  ciliatum    (L.)    Raf.,   358, 

4oo,  558,  589 

lanceolata  (Walt.)  A.  Gray,  589 
Stephanodiscus,  610,  614 

niagarse  Ehr.,  613 
Sterculia  platanifolia  Linn.,  79 
Sterculiaceas,  79 
Stereocaulon,  220,  236 

condensatum  Hoffm.,  16 

coralloides  Fr.,  229,  261 

paschale   (L.)   Fr.,   222,   224,   233, 

261,  278,  320,  662,  696 
Sticta,  236 

amplissima  (Scop.)  Mass.,  250 
crocata  (L. )  Ach.,  250 
fuliginosa,  250 
limbata  (Sm.)  Ach.,  229,  250 
pulmonaria    (L-)    Ach.,   250,  678, 

688 
scorbiculata    (Scop.)    Ach.,    232, 

251 
Stigeocloniutn,  26,  617 


INDEX. 


777 


Strophostyles    helvola    (L. )     Britton, 

394,  427,  541 
Styracaccie,  82 
Symphoricarpos  occideiitalis  Hook., 

430,  593 

racemosa  Michx.,  429 
symphoricarpos  (L. )  MacM.,  429, 

593 

Syuchytrium,  196 
Syndesmon     thalictroides      ( L. ) 

Hoffmg.,  363,  387 
Synedra,  6n 

pulchella  (Ralfs.)  Kg.,  613,  614 

ulna  (Nitzsch. )  Ehr.,  613,  614 

Syntherisma  sanguinalis    (L.)   Nash, 

374 
Syringa  vulgaris  L.,  430 

Tabcllaria  fenestrata,  614 

fenestrata  Lyndb.   Kg.  vnr.  inter- 
media Griin,  613 

flocculosa,  614 
Tagetes,  89 
Tanacetum,  545 

vulgare  L.,  414 
Taraxacum    taraxacum    (L. )     Karst., 

409,  424,  548,  593 
Taxaceae,  373 

Taxus  minor  (Michx.)  Britton,  373 
Tecoma,  126 

radicaus    (Linn.)   DC.,  82,  85,  87, 

124,  130,  131,  132,  133 
Tetraphis  pellucida  Hedw.,  42 
Tetragonanthusdeflexus  (J.  E.  Smith) 

Kuntze,  551 
Tetraspora,  617 

cylindrica  (Wahlenb. )  Ag.,  26 
Teucrium  canadense  L.,  358,  403,  590 
Thalesia  uniflora,  369 
Thalictrum  L.,  509,  550 

adiantifolium  Bess.,  513 

adriantoides  Hort.,  513 

alpinum  Linn.,  512 

aquilegifolium  Linn.,  514,  515 

ceesium  Greene,  517 

campestre  Greene,  516 

Carol i n ianu)n  Bosc.,  515 

caulophylloides  Small,  516 

clavatum  DC.,  512 

clavatum  Hook.,  512 

coriaceum  Small,  516 


Thalictrum 

Cornuti  Torr.  &  Gray,  514,  515 
var.  a  Hook.,  513 
var.  /?  Hook.,  514 
macrostylum  Shuttle.,  514 
var.  monostyla  Bot.  Zeit.,  514 
corynellum  DC.,  514 
Cuernavacanum  Rose,  518 
dasycarpum  Fisch.  &  Lall.,  513 
debile  Buckley,  515 

var.  Texanum  Gray,  515 
dioicum  L.,  363,  388,  515,  555 
var.  coriaceum  Britton,  516 
var.  oxycarpum  Torr.,  516 
Fendleri  Brew.  &  Wats.,  517 
Fendleri  Engelm.,  517 
Fendleri  I.  M.  Macoun,  516 

var.     platycarpum    Trelease, 

517 

var.  polycarpum  Torr.,  517 
var.  Wrightii  Trelease,  517 

filipes  Torr.  &  Gray,  512 

Galeottii  Lee.,  520 

gibbosum  Lee.,  520 

glaucum  Desf.,  513 

grandiflorum  Rose,  519 

grandifolium  Rose,  519 

grandifolium  Wats.,  522 

Guatemalense  Rose  &  DC.,  519 

Hernandezii  Tausch.,  520 

Hernandezii,  521 

hesperium  Greene,  517 

Jaliscanum  Rose,  518 

Kemense  Fries,  513 

lavigatum  Michx.,  515 

lanatum  Lcc.,  520,  521 

leucostfinon  Koch.  &  Bauch£,  514 

longist \lutn  DC.,  519 

macrostylum  Small  &  Heller,  514 

Madrense  Rose,  521 

minus  Linn.,  513 

var.  Kemense  Trelease,  513 
var.  adiantifolium  Hort.,  5f3 
var.  elatum  Lee.,  513 

nudicaule  Schweinitz,  512 

occidentale  Gray,  516 

Pachucense  Rose,  519 

papillosum  Rose,  521 

peltatum  DC.,  517 

petaloideum  Linn.,  512 

pinnatum  Wats.,  521 


778 


INDEX. 


Thalictrum 

platycarpum  Greene,  517 
polycarpum  Watson,  517 
polygamum  Muhl.,  514 

var.   macrostylum   Robinson, 

5H 
Pringlei  Wats.,  518 

var.  reticulaturn  Rose,  518 
pubigerum  Benth.,  519 
pubigerum  Pringle,  518 
purpurascens    L-,    363,    388,    427, 

5r3.  555,  582 

var.  ceriferum  Austin,  514 
purprtreum  Hort.,  513 
purpureum  Schang.,  513 
revolutum  DC.,  513 
revolutum  Lee.,  514 
Richardsonii  Gray,  512 
rugosum  Pursh,  513 
saxatile  Vill.,  513 
saxatilis  Hort.,  513 
sparciflorum  Turcz.,  512 
speciosum  Hort.,  513 
tomentellum   Robinson   &  Seat., 

52i 

venulosum  Trelease,  516 
Wrightii  Gray,  517,  522 
Thelocarpon    prasinellum     Nyl.,     18, 

228,  274,  306,  327 
Theloschistes,  672 

concolor  (Dicks.)  Tuck.,  13,  238, 

302,  304,  309,  668,  670,  685 
var.    effusa   Tuck.,   310,    670, 

685 
chrysopthaltnus  (L.)  Norm.,  302, 

305,  309,  670,  675,  685 
lychneus    (Nyl.)    Tuck.,  13,    244, 

292,  300,  302,  309,  666,  670,  685 
polycarpus    (Ehrh.)    Tuck.,    244, 

302,  304,  309,  670,  671,  685 
Thlaspi  arvense  L.,  191,  582 
Thuidium,  36 

abietinum  Sch.,  49 
philiberti  Limpr.,  49 
recognitum  Lindb.,  49 
Thuja  occidentalis  L.,  225 
Thymeleacese,  398 
Tilia,  429,  565,  566 

americana  L.,  363,  366,  396,  428, 

43°,  586 
Tiliaceae,  396,  586 


Toxylon   pomiferum  Raf.,  71,  84,  87, 

93,  130,  131,  132,  133 
Tradescantia,  54 

reflexa  Raf.,  377 

virginiana  L,.,  377 

Triadenum  virginicum  (L. )  Raf.,  586 
Trifolium  hybridum  L,.,  393 

pratense,  622,  627 

repens  L.,  393,  541,  622 
Triglochin  maritima  L,.,  572 
Trillium  cernuum  L.,  363,  378 

erectum  L,.,  363,  378 
Triosteum  perfoliatum  L.,  363,  408 
Triphragmium  Link.,  537,  554 

clavellosum  Berk.,  554 
Triticum  vulgare  L.,  548 
Typha  latifolia  L-,  373,  626,  624 
Typhacese,  373 

Uimacese,  70,  89,  383 
Ulmus,  85,  324 

americana  L-,  70,  87,  89,  130,  132, 

!33.  358,  363,  383 
fulva  Michx.,  70,  363,  383,  629 
racemosa  Thomas,  383 
Ulota  crispa  Brid.,  41 
curvifolia  Brid.,  41 
hutchinsiae  Sch.,  41 
Ulotrichiaceae  (Kg.)  Borzi,  26 
Umbelliferse,  89,  399,  588 
Umbilicaria,    4,    5,    6,    221,    236,     282, 

662 

dillenii  Tuck.,  3,  14,  249 
hyperborea  Hoffm.,  228,  250 
muhlenbergii  (Ach.)  Tuck.,   222, 

224,  249 
pustulata   (L. )  Hoffm.   var.  papu- 

losa  Tuck.,  224,  250 
vellea  (L.)  Nyl.,  233,  249 
Uncinula,  424 

clintonii  Peck,  428 
necator  (Schw.)  Burrill,  429 
salicis  (DC.  )  Wint.,  428 
Unifolium  canadense  (Desf. )  Greene, 

378,  577 
Urceolaria,  657 

actinostoma  Pers.,   281,  282,   287, 

288,  320 

scruposa  (L.)  Nyl.,  16,  223,  224, 
263,  286,  293,  296,  301,  320,  668, 
696 


INI)  I   \. 


779 


Uredo,  537 

agrimoniie-eupatorue  ( DC. )  Wint. , 

560 

alpcstris  Schrcet.,  640 
polypodii  (P.)  DC.,  560 
pyrola  (Gm.)  Wint.,  538 
Viola  Schum.,  633 
Uromyces  L/ink.,  537,  540 
albus  Diet.  &  Holw.,  541 
appendiculatus  (P.)  L,iuk.,  541 
argophyllse  Seym.,  543 
caladii  (Schw.)  Farlow,  542 
caryophyllinus  (Schrank.) 

Schroet.,  543 

euphorbiie  Cooke  &  Peck,  541 
fabie  ( P. )  De  Bary,  540,  554 
gerauii  (DC.)  Wint.,  556 
hedysari-paniculata   (Schw.)  Far- 
low,  543 

lespedezie  (Schw.)  Pk.,  543 
orobi  (P.)  Wint.,  541 
polygoni  (P.)  Fckl.,  542 
polymorphus  Pk.,  540 
pyriformis  Cooke,  543 
rudbeckise  Arth.  &  Holw.,  543 
terebinthi  (DC.)  Wint.,  543 
trifolii  (Alb.  &  Schw.)  Wint.,  541 
Uropyxis  Schroeter,  537,  544,  552 

amorphre  (Curt.)  Schroet.,  552 
Urtica,  546 

gracilis   Ait.,   358,   363,  383,    547, 

578,  626 

Urticacese,  383,  578 
Urticastrum  divaricatum  (L.)  Kuntze 

358,  363,  384,  578 
Utricularia  vulgaris  L,.,  355,  406 
Usnea,  225,  659,  676 

barbata  (L.)  Fr.,  684 

var.  ceralina  Schaer.,  243,  684 
var.    dasypoga   Fr.,    228,  243, 

665,  684 
var.   florida  Fr.,    13,  242,   309, 

670,  675,  677,  684 
var.  hirta  Fr.,  242,  301,  309 
var.  plicata  Fr.,  243 
var.  rubiginea  Michx.,  13,  301, 

309 
cavernosa    Tuck.,    227,    243,    662, 

677,  684 

longissima  Ach.,  219,  243 
trichodea  Ach.,  243 


Usnei,  675 

Ustulina   maxima   (Haller)    Schroter, 
564 

vulgaris  Tul  ,  564 
Uvularia  grandi flora,  363 

grandiflora  Sm.,  554 

grandiflora  J.  E.  Smith,  363,378,554 

perfoliata  L.,  554 

sessilifolia  L.,  554 

Vaccaria  vaccaria  (L. )  Britton,  581 

Vacciniaccte,  589 

Vaccinium  vitis-idsea  L.,  540 

Vaginariete  Gomont,  28 

Vagnera  racemosa   (L. )  Morong,  363, 

378,  577 
stellata  (L. )  Morong,  378,  577 

Valeriaua  edulis  Nutt,  367,  408 

Valerianacese,  408 

Vallisneriaceae,  374,  572 

Verbascum  thapsus  L-,  405 

Verbena,  426 

bracteosa  Michx.,  403 
hastata  L,.,  403,  426,  590,  626 
stricta  Vent.,  403,  426,  556 
urticifolia  I/.,  403,  426,  626 

Verbenaceae,  403,  590 

Verbesina  alternifolia,  23 

Vernonia  fasciculata  Michx.,  357,  410, 

594 

Veronica    americana    Schwein.,    360, 
406,  592 

peregrina  L-,  406 

scutellata  L.,  592 
Verrucaria,  670 

conoidea  Fr.,  708 

epigsea  (Peru.)  Ach.,  231,  275 

fuseella  Fr.,  18,  300,  328,  708 

muralis  Ach.,    18,    297,   300,  328, 
666,  667,  670,  708 

nigrescens    Pers.,    228,    275,    297, 
300,  328,  666,  670,  708 

viridula  Ach.,  666,  708 
Viburnum,  430 

dentatum  L,.,  407 

lentago  L.,  363,  4o8,  430,  593 

opulus  L.,  366,  407,  592 
Vicia,  55,  540 

americana   Muhl.,  541,  585 

faba,  56,  57,  58,  67 

linearis  (Nutt.)  Greene,  541 


780 


INDEX. 


Viola,  424,  631,  636 

adunca  Smith,  635,  639 
arenaria  DC.,  636     • 
blanda  Willd.,  544,  635 

amcena    (Le    Conte)    B.S.P., 

545 

palustriformis    A.    Gray,   545 
bi flora  L-,  640 
Canadensis  L.,  544,  587,  636 
cognata  Greene,  635,  637,  640 
cucullata  Ait.,  639 
delphinifolia  Nutt.,  639 
epipsila  L,ed.,  640 
glabella  Nutt.,  635,  638,  639 
hastata  L.,  636,  638 
Labradorica  Schrank.,  636 
lanceolata  L  ,  635,  638 
lobata  Benth.,  640 
mirabilis  L-,  640 
Montaneusis  Rydb.,  636,  639 
Nuttallii  Pursh,  632,  640 
obliqua  Hill,    361,    363,  397,  587, 

632,  633,  635 
ocellata    Torr.    &   Gr.,    636,    639, 

640 

ovata  Nutt.,  632 
palustris  L,.,  640 

pedata  L.,  367,  397,  587,  632,  633 
pedatifida  Don. ,  367,  397,  632 
primulsefolia,  632,  635,  638 
pubescens  Ait.,  363,  398,  636,  637 
renifolia  A.  Gray,  635 
rostrata  Pursh,  636 
rotundifolia  Michx.,  635 
sagittata  I/.,  632,  633 
scabriuscula  (T.  &  G.)  Schw.,  545, 

636 


Viola 

septentrionalis  Greene,  635 

striata  Ait.,  632,  636 

tricolor  I,.,  632 
Violaceae,  397,  587 
Viscum  orientale  Willd.,  170 
Vitacese,  78,  119,  396,  586 
Vitis,  120 

cordifolia  Michx.,  78,  84,  87,  119, 
131,  132,  133 

vulpina  L,.,  358,363,  396,  586 
Volvox  globata,  617 

Washingtonia  claytoni  (Michx.)  Brit- 
ton,  363,  399,  544 

longistylis  (Torr.)  Britton,  588 
Webera  nutans  Hedw.,  42 
Woodsia  ilvensis  (L,. )  R.  Br.,  614 

oregana  D.  C.  Eaton,  369,  370 

Xanthium,  551 

canadense    Mill.,    409,    551,    594, 

626 
Xanthoxylum  americanum  Mill.,  363, 

394,  585 
Xylaria,  564 

clavata  (Scop.)  Schranck,  567 
polymorpha  (Pers.)  Greville,  567 
Xylariacese,  563 

Zea  mays  L,.,  55,  58,  65,  08,  550 
Zizania  aquatica  L.,  357,  375.  573,  6a6, 

650 

Zizia  aurea  (L. )  Koch,  361,  399,  588 
cordata  (Walt.)  Koch,  399,  588 
Zygadenus  elegans  Pursh,  367,  377 
Zygnemacese  (Menegh.)  Rabenh.,  26 


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